Rosids: Fabids Spring 2012

Fig. 8.1

Rosids – Major Points Comprise about 25% of all angiosperms Includes two main clades: fabids and malvids Main support for monophyly from molecular data No clear morphological synapomorphies, but tendencies to have perianths with unfused parts and a stamen merosity > calyx or corolla, although there are many exceptions Extreme variation in habit (trees, shrubs, herbs, vines, etc.) as well as extensive proliferation of floral syndromes, including wind, insect, bird, and bat pollination Transition from apocarpy to syncarpy as seen before; fusion and embellishment of floral parts

Fig. 8.30

Core Eudicots: The Rosids Fabids: Order Malpighiales Order Fabales Order Rosales Order Cucurbitales Order Fagales Malvids: Order Myrtales Order Brassicales Order Malvales Order Sapindales

Core Eudicots: The Rosids - Fabids Order Malpighiales Euphorbiaceae* – Spurges Salicaceae* – Willows and poplars Violaceae – Violets Order Fabales Fabaceae* – Beans Order Rosales Rosaceae* – Roses Moraceae – Figs, mulberries Ulmaceae – Elms Order Cucurbitales Curcurbitaceae* – Cucumbers, squashes Begoniaceae – Begonias Order Fagales Betulaceae – Birches Fagaceae – Oaks, beeches, chestnuts Juglandaceae – Walnuts, hickories *Family required for recognition

Rosids-Fabids: Malpighiales: Euphorbiaceae (The Spurge Family) Widespread, but most diverse in tropical regions Trees, shrubs, herbs, or vines, sometimes succulent; leaves usually alternate Diversity: ca. 6,300 species in 218-245 genera Flowers: Unisexual; sepals 2-6; petals 0-5; carpels usually 3, ovule 1 per locule; styles usually 3 and sometimes divided; inflorescences often highly modified; fruit a schizocarp, seeds usually arillate Significant features: Often with latex/laticifers (toxic) Special uses: rubber (Hevea), cassava/manioc (Manihot), poinsettia (Euphorbia), ornamentals Required taxa: Euphorbia

Euphorbiaceae: Euphorbia Ca. 2,400 species White latex (usually) One female and few to many male flowers aggregated into a cyathium (one type of false flower or pseudanthium) Cyathium subtended by modified leaves (cyathophylls)

Euphorbiaceae: Euphorbia cyathium From the Euphorbia PBI website

Euphorbiaceae: Euphorbia Digital Flowers

castor bean ~ poisonous seeds Euphorbiaceae Castor oil plant (Ricinus communis) castor bean ~ poisonous seeds ^

Euphorbiaceae Rubber (Hevea brasiliensis) ^

Euphorbiaceae ^ Tapioca, Cassava (Manihot) Manihot esculenta cassava, tapioca ^

Euphorbiaceae Tung oil (Aleurites) Aleurites fordii tung-oil tree ^

Rosids-Fabids: Malpighiales: Salicaceae (The Willow or Poplar Family) Widespread, from tropical to north temperate and boreal regions Trees or shrubs Diversity: 1,200 species in 54-55 genera Flowers: bisexual or unisexual; sepals 3-8; petals 3-8; stamens 2-∞; carpels 2-4, connate, in superior ovary; fruit variable Significant features: leaves simple, often with salicoid teeth; salicin in most; includes what was formerly called the “Flacourtiaceae” Special uses: lumber, shade trees, ornamentals Required taxa: Populus, Salix

Salicaceae: Salix -bud scale single -catkins usually erect or ascending -flowers unisexual -each flower with 1-4 basal nectar glands -stamens 1-12 -mainly insect pollination

Salicaceae: Populus -bud scales several, overlapping -catkins arching or drooping -flowers unisexual -each flower with a basal cup-like disk -stamens 8-numerous -wind-pollination

Rosids-Fabids: Malpighiales: Violaceae (The Violet Family) Widespread, but predominantly herbs of temperate regions Herbs, shrubs, or trees Diversity: 700-800 species in 23-24 genera Flowers: Sepals 5; petals 5; 5 connivent stamens; carpels usually 3, connate, superior ovary; fruit usually a loculicidal capsule Significant features: Zygomorphy, nectar spurs; floral cleistogamy Special uses: Violets grown primarily as ornamentals Family not required

Violaceae: connivent stamens

Violaceae: Viola -mostly herbs, some shrubs -flowers zygomorphic -lower petal spurred -spring flowers open-pollinated, summer flowers remaining closed (cleistogamous)

Rosids-Fabids: Fabales: Fabaceae (The Legume Family) Nearly cosmopolitan Herbs, vines, trees, shrubs with usually alternate, stipulate, pinnately to palmately compound leaves (sometimes unifoliolate or simple) Diversity: 19,500 species, 720-730 genera – THIRD LARGEST FAMILY of angiosperms Flowers: a short, cup-like hypanthium present; sepals & petals usually 5, free or connate; petals all alike or the uppermost 1 differentiated (banner), the lower 2 forming a keel or flaring apart; stamens 5 or 10-many, if connate then monadelphous or diadelphous; carpel 1, on a short stalk (gynophore); fruit is a legume (Duh!) but sometimes modified Significant features: High nitrogen metabolism w/ unusual amino acids, often with root nodules with N-fixing bacteria; leaf and leaflet pulvinuses well developed; endosperm often lacking; wide range of floral diversity; 3 subfamilies but 1 is not monophyletic Special uses: Many!! Beans, peas, peanuts, soybean, clover, ornamentals (Mimosa, Bauhinia); lumber, dyes, resins Required taxa: Glycine, Trifolium, Mimosa, Cercis, Gleditsia and the three subfamilies

Fabaceae vegetative characters root nodules pulvinus compound leaves

Fabaceae floral characters Diadelphous stamens: 9 + 1 Perigynous flower, short hypanthium gynophore Marginal (parietal) placentation

Fabaceae fruit and seed characters non-endospermous seeds legumes, loments, etc.

Mimosoideae “Caesalpinioideae” Papilionoideae Leaves pinnately Leaves usually twice pinnately compound Fls actinomorphic, petals valvate, distinct or basally fused Stamens 10-many, “Caesalpinioideae” Leaves usually pinnately or twice pinnately compound Fls + weakly zygomorphic, upper petal usually innermost; petals distinct Stamens 5 or 10, distinct Papilionoideae Leaves pinnately compound to trifoliolate Fls zygomorphic, upper petal (banner) outermost; well defined wings and keel Stamens 10, monadelphous or diadelphous

Fabaceae – Subfamily Mimosoideae Albizia julibrissin Acacia sp.

Fabaceae: Mimosoideae Actinomorphic tubular flowers in heads many stamens, not fused Albizia julibrissin mimosa, silktree

Fabaceae: Mimosoideae: Mimosa -woody or herbaceous -often armed (with prickles) -leafstalk without glands -flowers in heads or rarely racemes or spikes -stamens 10 or fewer

Fabaceae – Subfamily “Caesalpinioideae”

Fabaceae: “Caesalpinioideae” zygomorphic flower pulvinus Stamens not fused -10 or fewer Senna obtusifolia sicklepod

Fabaceae: “Caesalpinioideae”: Cercis -unarmed -leaves simple, palmately veined -flowers clustered, appearing before leaves -corolla rose to pink-purple Cercis canadensis - redbud

Fabaceae: “Caesalpinioideae”: Gleditsia -armed (with thorns) -leaves 1- or 2-pinnate -flowers small, unisexual or bisexual -staminate inflorescences catkin-like, pendent -fertile inflorescences with bisexual or carpellate flowers Honey locust

Fabaceae – Subfamily Faboideae

Fabaceae: Faboideae Petals unequal: banner wings keel monadelphous stamens Crotalaria spectabilis showy rattlebox bacterial root nodule

Crotalaria spectabilis Fabaceae: Faboideae Petals unequal: banner wings keel Crotalaria spectabilis showy rattlebox

Fabaceae: Faboideae Lathyrus sweet-pea Digital Flowers

Fabaceae: Faboideae: Glycine -leaves pinnately 3-foliolate -inflorescence a raceme -stamens diadelphous -seeds few per pod

Fabaceae: Faboideae: Trifolium -leaves palmately (or pinnately) foliolate with usually 3 leaflets -inflorescences racemose but often appearing head-like -stamens diadelphous -fruits enclosed by the persistent corolla -seeds 1-6 per pod

Rosids-Fabids: Rosales: Rosaceae (The Rose Family) Cosmopolitan, primarily in the Northern Hemisphere Herbs, shrubs or trees (75% woody plants) Diversity: 2,500-3,000 species in 85-90 genera Flowers: Showy, actinomorphic, hypanthium present; sepals 5; petals 5; stamens usually numerous; carpels 1 to many, apocarpous or syncarpous; ovary superior or inferior; fruit can be a follicle, achene, pome, drupe, or associated with expanded receptacle Significant features: Wide range of fruit evolution within family; leaves alternate, stipules present Special uses: Fruits (apples, pears, berries), ornamental herbs, trees, and shrubs; lumber, perfumery Required taxa: Rubus, Prunus

Rosaceae: Rosa -shrubs, often prickly -leaves pinnately compound -ovary superior -hypanthium + globose and fleshy, contracted at the mouth -carpels numerous -fruit an achene

Rosaceae: Rubus* -herbaceous to shrubby, usually with prickles -leaves often compound with 3 to 7 leaflets -carpels usually numerous, borne on an elongate receptacle -fruit a drupelet, forming an aggregate fruit (blackberries and raspberries) *required for recognition

Rosaceae: Prunus* -trees or shrubs -bark with prominent horizontal lenticels -ovary superior -carpel solitary -fruit a drupe, endocarp stony *required for recognition

Rosaceae: Malus -trees with simple leaves -ovary inferior, with 5 carpels -mature carpels papery or softly leathery -fruit a pome, lacks stone cells

Rosids-Fabids: Rosales: Moraceae (The Fig Family) Widespread, from tropical to temperate regions Trees, shrubs, or vines (sometimes herbs) Diversity: 1,500 species in 53 genera Flowers: Unisexual, inconspicuous; tepals 0-4 or 5 (-8); carpels usually 2, connate, superior ovary; inflorescences cymose, highly modified, compact, receptacle expanded; fruit is a drupe, often in a multiple fruit structure (syconium). Significant features: laticifers/latex throughout the plant Special uses: figs (Ficus), mulberries (Morus), breadfruit (Artocarpus), ornamentals, e.g. osage orange (Maclura) Family not required

Moraceae Dorstenia Ficus carica – Cultivated Fig Artocarpus (breadfruit) Maclura pomifera Osage orange Morus rubra - Mulberry

Moraceae: Ficus -shrubs or trees -connate stipules enclosing the terminal buds -leaves with entire margins -flowers minute, borne inside the syconium -wasp-pollinated

Moraceae – The Fig and The Fig Wasp

Rosids-Fabids: Rosales: Ulmaceae (The Elm Family) Widely distributed; maximal diversity in temperate regions of N. Hemisphere Trees with alternate, 2-ranked leaves Diversity: 35 species in 6 genera Flowers: Small, inconspicuous; tepals 4-9; stamens 4-9; carpels 2, connate, superior ovary; fruit a samara or nutlet, seeds flat Significant features: Leaves simple with pinnate venation, margins simply or doubly serrate, blade base asymmetrical; endosperm of a single layer Special uses: Elms provide lumber; some trees used as ornamentals Family not required

Ulmaceae: Ulmus

Rosids-Fabids: Cucurbitales: Cucurbitaceae (The Cucumber or Squash Family) Widespread in the tropics and subtropics, a few in temperate regions Herbaceous or soft woody vines with scabrous stems and palmately veined/lobed leaves and usually with tendrils Diversity: 900 species in 118-122 genera Flowers: hypanthium present; sepals & petals 5, usually connate; stamens 3-5; carpels usually 3; ovary half-inferior or inferior; fruit usually a berry (with hardened rind a pepo); seeds flattened, the seed coat with several layers Significant features: wide range of floral diversity, “toothed” leaves lacking stipules; female flowers epiperigynous Special uses: cucumbers (Cucumis), pumpkins, gourds, and squashes (Cucurbita), watermelons (Citrullus) etc. are eaten for fruits and seeds; Luffa, some ornamentals Required taxa: family only

Cucurbitaceae: Cucurbita -trailing herbs -leaves large, cordate- angled or lobed -flowers large, solitary in axils -corolla campanulate, deeply 5-lobed -ovaries and fruits smooth or hairy, not prickly -fruits large, with a firm rind -gourds, squashes, pumpkin

Rosids-Fabids: Cucurbitales: Begoniaceae (The Begonia Family) Widely distributed in the tropics and subtropics Herbs or soft woody shrubs Diversity: ca. 1,400 species in 2 genera (all but one species in Begonia) Flowers: Unisexual; tepals petaloid, 2-10 but usually 4 in 2 whorls (staminate) and 5 in 1 whorl (pistillate); carpels usually 3, connate; inferior ovary; fruit a loculicidal capsule, usually winged Significant features: Soft herbs, typically of shaded habitats; stigmas elongated, twisted, yellow, papillose Special uses: primarily ornamentals Family not required

Begoniaceae: Begonia -winged ovaries Carpellate Flowers Staminate Flowers -winged ovaries

Rosids-Fabids: Fagales: Fagaceae (The Oak and Beech Family) Widespread, in tropical to temperate regions of the Northern Hemisphere Trees and shrubs Diversity: 670-970 species in 7 genera Flowers: Unisexual (monoecious); tepals usually 6 and reduced, inconspicuous; stamens 4-many; carpels 3 (-12), connate, inferior ovary; fruit a nut, associated with a spiny or scaly cupule Significant features: Male inflorescences in dangling catkins; female inflorescences in sessile clusters Special uses: edible nuts (chestnuts), lumber, tannin, cork; ornamental trees Family not required

Fagaceae: Quercus Bark pale to dark but scaly or furrowed Buds clustered at twig tips, ovate Leaves lobed or unlobed Male flowers in drooping catkins Cupule saucer-like or cup-shaped Nut circular in cross-section

Fagaceae: Fagus Bark light gray, smooth Buds solitary at twig tips, slender and acute Leaves unlobed, strongly straight-veined Male flowers in a rounded head Cupule with 4 valves Nut compressed or triangular

Rosids-Fabids: Fagales: Betulaceae (The Birch Family) Widespread, in temperate to boreal regions, primarily of the northern hemisphere Trees or shrubs; leaves doubly serrate Diversity: 140 species in 6 genera Flowers: Unisexual (monoecious); tepals (0-) 1-4 (-6), highly reduced; stamens 1-4; carpels 2, connate, inferior ovary; fruit an achene, nut or 2-winged samara Significant features: Flowers in erect (female) or pendant (male) catkins (aments); staminate and carpellate flowers in separate inflorescences Special uses: hazel nuts edible; lumber, shade trees, ornamentals Family not required

Betulaceae: Betula Outer bark often separating in thin sheets Carpellate and staminate flowers both in bracteate catkins Carpellate catkins ovoid to cylindrical, with 2-3 flowers per bract and the bracts papery (bracteate dichasia) Staminate flowers 3 per bract; stamens 2, bifid

Rosids-Fabids: Fagales: Juglandaceae (The Walnut and Hickory Family) Widespread from tropical to temperate regions Aromatic trees; leaves pinnately compound or trifoliolate, usually alternate and spiral Diversity: 59 species in 8 genera Flowers: Unisexual (monoecious or dioecious); tepals 0-4, inconspicuous; stamens 3-many; carpels usually 2, connate, ovary inferior; fruit a nut or nutlet Significant features: Fruit often associated with bracts or bracteoles that form an outer “husk” Special uses: fruits of hickories (Carya) and walnuts (Juglans) are eaten; walnut and hickory are valued for their lumber; some ornamentals Family not required

Juglandaceae: Juglans Twigs with chambered pith Leaflets all about the same or the median ones largest Staminate catkins sessile, solitary Nut with an indehiscent, usually rough or furrowed husk

Juglandaceae: Carya Twigs with solid pith Apical leaflets largest Staminate catkins sessile or pedunculate, in clusters Nut with a dehiscent or partially dehiscent, often smooth husk