IB335 - Lab Manual - Spring 2015 PDF

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INTEGRATIVE BIOLOGY 335 SPRING 2015

SYSTEMATICS
OF PLANTS

FAMILY NOTES & LABORATORY EXERCISES

MAGNOLIID CLADE

MONOCOT CLADE

BASAL EUDICOTS

CARYOPHYLLALES

ROSID CLADE

ASTERID CLADE

STEPHEN R. DOWNIE ▪ DANIELLE M. RUFFATTO ▪ KENNETH R. ROBERTSON


Dear Botanist-In-Training,

From Charles Darwin to John Audubon and Aldo Leopold, the naturalists of old were extraordinary
observers. Further, they could capture all the natural diversity they saw with written descriptions
and detailed drawings. Though this class was intended to give you an overview of approximately
40 of the most widespread and/or economically important angiosperm families and subfamilies, it is
also meant to teach you how to really observe and describe plants.

Trained as a scientist and artist, I try to view the world around me through the eyes of the early
naturalists. So, when I sat down to design this new version of the IB 335 Systematics of Plants
Lab Manual, I thought of Georgia O’Keeffe’s quote, “When you take a flower in your hand and
really look at it, it’s your world for the moment. I want to give that world to someone else.” Thus, I
designed each lab exercise so that you will not only be examining the flowers, leaves, stems and
roots of plants, but also describing your observations through answering questions and creating
simplistic diagrams.

Hopefully by the end of the semester all of you will recognize the “beauty in the details.”

Best wishes,

Danielle M. Ruffatto
INTEGRATIVE BIOLOGY 335
table of contents

IB 335 Plant Families 5 Orchidaceae46


Iridaceae48
Classification of Plant Families 5 Liliaceae49
Magnoliaceae6 Poaceae50
Ranunculaceae7
Papaveraceae8
Cactaceae9 Laboratory 1 53
Caryophyllaceae10
Woody Twig Terminology Lab Exercise 53
Portulacaceae11
Dichotomous Key Guidelines 56
Polygonaceae12
Dichotomous Key for Woody Plants 57
Hamamelidaceae13
Glossary of Woody Twig Terminology 59
Moraceae14
Fagaceae15
Betulaceae16 Laboratory 2 63
Cucurbitaceae17
Violaceae18 Vegetative Terminology Lab Exercise 64
Salicaceae19 Glossary of Vegetative Terminology 76
Fabaceae20
Mimosoideae21 Laboratory 3 83
Caesalpinioideae22
Faboideae23 Floral Drawing Guidelines 83
Onagraceae24 Floral Formula Guidelines 84
Euphorbiaceae25 Floral Terminology Lab Exercise 88
Rosaceae26 Inflorescence Terminology Lab Exercise 94
Spiraeoideae27 Glossary of Floral Terminology 97
Rosoideae28 Glossary of Inflorescence Terminology 104
Amygdaloideae29
Maloideae30
Laboratory 4 109
Brassicaceae31
Malvaceae32 Fruit Terminology Lab Exercise 109
Aceraceae33 Morphology-Based Fruit Types 110
Ericaceae34 Taxonomy-Based Fruit Types 117
Asclepiadaceae35 Special Fruit Types 121
Solanaceae36 Glossary of Fruit Terminology 123
Oleaceae37
Scrophulariaceae38
Lamiaceae39 Laboratory 5 127
Caprifoliaceae40
Magnoliaceae129
Asteraceae41
Ranunculaceae133
Apiaceae43
Papaveraceae137
Araceae44
Arecaceae45
Laboratory 6 141 Iridaceae238
Liliaceae240
Hamamelidaceae142 Arecaceae242
Moraceae144 Poaceae244
Fagaceae147
Betulaceae150
Practice Lecture Exam 1 247

Laboratory 7 153
Practice Lecture Exam 2 255
Cactaceae154
Caryophyllaceae157
The Story of Blackeyed Sue 265
Portulacaceae160
Polygonaceae162

Laboratory 8 165

Cucurbitaceae167
Violaceae170
Salicaceae172
Brassicaceae174
Malvaceae176
Ericaceae179

Laboratory 9 181

Fabaceae182
Onagraceae189
Euphorbiaceae191

Laboratory 10 193

Asclepiadaceae194
Solanaceae197
Oleaceae199
Caprifoliaceae202

Laboratory 11 205

Scrophulariaceae206
Lamiaceae208
Asteraceae211

Laboratory 12 217

Rosaceae219
Aceraceae227
Apiaceae230

Laboratory 13 233

Araceae234
Orchidaceae236
INTEGRATIVE BIOLOGY 335
CLASSIFICATION OF PLANT FAMILIES

The following classification of the plant


EUDICOT (TRICOLPATE) CLADE (CONTD.)
families covered in Integrative Biology 335 is
summarized from Judd et al. (2008, Table 9.1).
ROSID CLADE (ROSIDS)

EUROSIDS I (FABIDS)
MAGNOLIID CLADE (MAGNOLIIDS) Moraceae
Fagaceae
Magnoliaceae Betulaceae
Cucurbitaceae
Violaceae
Salicaceae
MONOCOT CLADE Fabaceae
Onagraceae
Araceae Euphorbiaceae
Arecaceae Rosaceae
Orchidaceae
Iridaceae
EUROSIDS II (MALVIDS)
Liliaceae
Poaceae Brassicaceae
Malvaceae
Aceraceae

EUDICOT (TRICOLPATE) CLADE


ASTERID CLADE (ASTERIDS)
BASAL EUDICOT (BASAL TRICOLPATE) CLADE Ericaceae
Ranunculaceae
Papaveraceae EUASTERIDS I (LAMIIDS)
Asclepiadaceae
Solanaceae
CORE EUDICOT (CORE TRICOLPATE) CLADE
Oleaceae
CARYOPHYLLALES Scrophulariaceae
Lamiaceae
Cactaceae
Caryophyllaceae
Portulacaceae EUASTERIDS II (CAMPANULIIDS)
Polygonaceae Caprifoliaceae
Asteraceae
Hamamelidaceae Apiaceae

IB 335 Plant Families 5


MAGNOLIACEAE
magnolia family ♂ Ca3(x) Co6(x) A∞ G∞

Magnoliid Complex (Magnoliid Clade)

FLORAL CHARACTERISTICS EXAMPLES


• Large perfect flowers Magnolia spp. [Magnolia]
• Actinomorphic symmetry M. acuminata (L.) L. [Cucumbertree]
• Numerous spirally arranged parts M. acuminata var. acuminata
• No adnation or connation M. acuminata var. subcordata (Spach) Dandy [Yellow
• Anthers LAMINAR Cucumbertree]

• Apocarpous gynoecium M. fraseri Walter [Fraser Magnolia]

• Superior ovary M. grandiflora L. [Southern Magnolia]

• Marginal placentation M. grandiflora × M. virginiana = Magnolia ‘Freeman’


M. macrophylla Michx. [Bigleaf Magnolia]
M. sieboldii K. Koch [Oyama Magnolia]
INFLORESCENCE TYPE(S) Magnolia × soulangiana Soul.-Bod. [Saucer Magnolia]
• Flowers solitary and terminal M. stellata (Siebold & Zucc.) Maxim. [Star Magnolia]
M. tripetala (L.) L. [Umbrella Magnolia]
M. virginiana L. [Sweet Bay]
FRUIT TYPE(S)
Liriodendron spp. [Tuliptree]
• An aggregate of follicles or samaras L. tulipifera L. [Tulip poplar]
• Seeds often surrounded by a fleshy aril L. tulipifera ‘Aureo-marginatum’
L. chinense (Hemsl.) Sarg. [Chinese tuliptree]

HABIT
• Trees or less often shrubs
• Aromatic with ethereal oil cells

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Entire margin or lobed at tip in Liriodendron
• Stipules and stipular scars surrounding
twigs

NOTE: Several interesting Magnolias are planted


in the southeastern corner of Mt. Hope Cemetery
[just south and west of the Meat Science Lab and
West of Florida Avenue Residence Halls] and also
at the corner of Nevada and Busey in Urbana.

6 Integrative Biology 335: Plant Systematics


RANUNCULACEAE
buttercup family ♂ Ca4-∞ Co0-∞ A∞ G∞

Eudicots: Basal Eudicots

FLORAL CHARACTERISTICS EXAMPLES


• Perfect or rarely imperfect flowers [then Aconitum columbianum Nutt. [Columbian Monkshood]
dioecious plants] Actaea spp. [Baneberry]
• Actinomorphic or zygomorphic symmetry A. pachypoda Elliott [White Baneberry]
• Parts spirally arranged A. rubra (Aiton) Willd. [Red Baneberry]
• Stamens NOT LAMINAR Anemone spp. [Anemone]
• Apocarpous gynoecium A. blanda Schott & Kotschy [Greek Thimbleweed]
• Superior ovary with 1 locule A. coronaria L. [Lilies-of-the-Field]
• Numerous marginal ovules or 1 basal A. cylindrica A. Gray [Thimbleweed]
ovule
A. patens L. [Pasque Flower]
• Many adaptations for pollination
Anemone x fulgens (DC.) J. Gay ex Rchb. [Windflower]
Aquilegia canadensis L. [Red Columbine]
INFLORESCENCE TYPE(S) Caltha palustris L. [Yellow Marsh Marigold]
Cimicifuga racemosa (L.) Nutt. [Black Cohosh]
• Various cymes, racemes or panicles
Clematis L. spp. [Virgin’s Bower]
• Terminal
Delphinium tricorne Michx. [Dwarf Larkspur]
Eranthis hyemalis (L.) Salisb. [Winter Aconite]
FRUIT TYPE(S) Hepatica nobilis Schreb. [Liverleaf]
• Follicles or achenes Ranunculus septentrionalis Poir. [Buttercup]
• Rarely berries from one carpel Thalictrum spp. [Meadow-rue]
• Frequent adaptations for dispersal T. revolutum DC. [Waxyleaf Meadow-rue]
T. thalictroides (L.) Eames & B. Boivin [Rue Anemone]

HABIT
• Perennial and annual herbs
• Rarely woody vines

LEAF CHARACTERISTICS
• Simple or compound leaves
• Alternate arrangement
• Without stipules

IB 335 Plant Families 7


PAPAVERACEAE
poppy family ♂ Ca2(3) Co4-8 A6 or ∞ G 2-∞

Eudicots: Basal Eudicots

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Argemone mexicana L. [Mexican Prickly Poppy]
• Actinomorphic symmetry [zygomorphic in Bocconia frutescens L. [Parrotweed]
Fumarioideae] Chelidonium majus L. [Celandine]
• Perianth parts distinct Eschscholzia californica Cham. [California Poppy]
• Sepals CAUDUCOUS Hunnemannia fumariifolia Sweet [Mexican Tulip Poppy]
• Petals sometimes crumpled in bud Macleaya cordata (Willd.) R. Br. [Plume Poppy]
• Numerous stamens [or only 6 in Meconopsis grandis Prain [Blue Poppy]
Fumarioideae]
Papaver spp. [Poppy]
• Syncarpous gynoecium
P. orientale L. [Oriental Poppy]
• Superior ovary
P. somniferum L. [Opium Poppy]
• Usually 1-locular ovary with many parietal
ovules Sanguinaria canadensis L. [Bloodroot]
Stylophorum diphyllum (Michx.) Nutt. [Celandine Poppy]

INFLORESCENCE TYPE(S)
FUMARIOIDEAE EXAMPLES
• Flowers solitary or in cymes and racemes
Corydalis spp. [e.g. Corydalis, Fumewort]
Dicentra spp. [Squirrel-corn]
FRUIT TYPE(S)
D. canadensis (Goldie) Walp. [Squirrel corn]
• Capsules [PORICIDAL in Papaver] D. cucullaria (L.) Bernh. [Dutchman’s breeches]
• Seeds often with fleshy ELAIOSOMES for D. spectabilis (L.) Lem. [Bleeding heart]
ant dispersal

HABIT
• Annual or perennial herbs, rarely woody
• Sap milky or colored [often bright orange,
red or yellow (clear in Fumarioideae)]
• Some with alkaloids

LEAF CHARACTERISTICS
• Alternate to opposite or whorled
arrangement
• Entire to lobed or dissected margin

8 Integrative Biology 335: Plant Systematics


CACTACEAE
cactus family ♂ Ca∞ Co∞ A∞

G 3-∞
Eudicots: Caryophyllid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Often large and showy perfect flowers Carnegia gigantea (Engelm.) Britton & Rose [Saguaro]
• Actinomorphic to somewhat zygomorphic Cereus spp. [Night-blooming Cactus]
symmetry Echinocereus spp. [Hedgehog Cactus]
• Gradual transition from sepals to petals, Epiphyllum spp. [Orchid Cactus]
both numerous
Lophophora williamsii (Lem. ex Salm-Dyck) J.M. Coult [Peyote]
• Syncarpous gynoecium made up of many
connate carpels Mammillaria spp. [Pincushion Cactus]

• Inferior ovary sunken into stem tissue Opuntia spp. [Pricklypears] = 3 found in IL:

• 1-locular ovary with many ovules O. fragilis (Nutt.) Haw. [Brittle Pricklypear: Endangered in IL]

• Parietal placentation O. humifusa (Raf.) Raf. [Devil’s-tongue]

• Sometimes a hypanthium present O. macrorhiza Engelm. [Twistspine Prickly Pear]


Rhipsalis baccifera (Sol. ex J.S. Muell.) Stearn [Mistletoe
Cactus]
INFLORESCENCE TYPE(S) Schlumbergera truncata (Haw.) Moran [False Christmas
Cactus]
• Solitary flowers emerging from AREOLES

FRUIT TYPE(S)
• Berries surrounded by stem tissue
• Seeds with embryo coiled around
perisperm

HABIT
• Perennial stem succulents, some tree-like
or epiphytic
• BETALAIN pigments
• SPINES produce from an AREOLE, which
sometimes contains numerous GLOCHIDS
• Native to the New World

LEAF CHARACTERISTICS
• Reduced or absent except in Pereskia

IB 335 Plant Families 9


CARYOPHYLLACEAE
pink family ♂ Ca 5 Co5 A10 G 2-5

Eudicots: Caryophyllid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Agrostemma spp. [Corncockle]
• Actinomorphic symmetry Dianthus spp. [Pinks, Carnations]
• Sepals connate and persistent Gypsophila spp. [Baby’s Breath]
• Petals distinct and often clawed at base Lychnis chalcedonica L. [Maltesecross]
and apically notched Saponaria officinalis L. [Bouncingbet]
• Syncarpous gynoecium often made up of 3 Silene spp. [Campion, Catchfly]
connate carpels
S. regia Sims [Royal Catchfly]
• Superior ovary
Stellaria spp. [Chickweed, Starwort]
• 1-locular ovary with many free central
ovules
• Possess an ANDROGYNOPHORE [or
GYNOPHORE] - a stalk that supports the
corolla, androecium and gynoecium

INFLORESCENCE TYPE(S)
• Basically cymes

FRUIT TYPE(S)
• Capsules with apical teeth [denticidal
capsules] or achenes in one group
• Seeds with embryo coiled around
perisperm

HABIT
• Annual or perennial herbs
• ANTHOCYANIN pigments
• Stems often with swollen nodes

LEAF CHARACTERISTICS
• Simple leaves
• Opposite arrangement
• Entire margin
• Leaf bases often connate and sheathing

10 Integrative Biology 335: Plant Systematics


PORTULACACEAE
purslane family ♂ Ca2 Co5-∞ A5-∞ G 2-9

Eudicots: Caryophyllid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Claytonia virginica L. [Virginia Spring Beauty]
• Actinomorphic symmetry Lewisia rediviva Pursh [Bitter Root]
• 2 persistent sepals [may be bracts] Portulaca grandiflora Hook. [Rose Moss]
• 5 to many petals [may be sepals] P. oleracea L. [Little Hogweed]
• 5 [then opposite petals] to many stamens Talinum spp. [Fameflower]
• Syncarpous gynoecium made up of 2-9 T. rugospermum Sims [Prairie Fameflower]
connate carpels Stellaria spp. [Chickweed, Starwort]
• Superior or rarely half-inferior ovary
• 1-locular ovary with 2 to many basal or free
central ovules

INFLORESCENCE TYPE(S)
• Flowers solitary or cymes, racemes or
heads

FRUIT TYPE(S)
• Capsules, sometimes circumscissile
capsules [PYXIS]
• Rarely a nut
• Seeds with embryo coiled around
perisperm

HABIT
• Annual or perennial herbs, less often
shrubs or small trees
• BETALAIN pigments

LEAF CHARACTERISTICS
• Simple leaves
• Alternate, opposite or basal arrangement
• Entire margin
• Leaves often succulent

IB 335 Plant Families 11


POLYGONACEAE
buckwheat family ♂ Ca3+3 (5) Co0 A3-9 G 2-3

Eudicots: Caryophyllid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect or rarely imperfect flowers Coccoloba uvifera (L.) L. [Seagrape]
• Actinomorphic symmetry Fagopyrum esculentum Moench [Buckwheat]
• Sepals often in 2 whorls of 3, distinct or Polygonum spp. [Knotweeds and Smartweeds]
connate at the base, sometimes an inner Rheum rhabarbarum L. [Garden Rhubarb]
sepal fused with an outer one
Rumex spp. [Docks]
• Petals absent
• Syncarpous gynoecium made up of 2-3
connate carpels
• Superior ovary
• 1-locular ovary with 1 basal ovule

INFLORESCENCE TYPE(S)
• Mostly racemes, spikes or panicles
• Sometimes in small axillary clusters

FRUIT TYPE(S)
• Triangular or lens-shaped achenes
• Embryo straight to curved

HABIT
• Annual or perennial herbs, sometimes
shrubs or even trees in the tropics
• ANTHOCYANIN pigments

LEAF CHARACTERISTICS
• Alternate, seldom opposite, whorled or
basal arrangement
• Entire margin
• Stipules usually present and sheathing
around stem [OCREA]

12 Integrative Biology 335: Plant Systematics


HAMAMELIDACEAE ♂ Ca 4-5 Co4-5(10) A4-5 OR

witch hazel family
G②
Eudicots: Core Eudicot ♂ Ca 4-5 Co4-5(10) A4-5 G②

FLORAL CHARACTERISTICS EXAMPLES


• Perfect or imperfect flowers [then Corylopsis spp. [Winter Hazel]
monoecious plants] Fothergilla gardenii L. [Dwarf Witch Alder]
• Actinomorphic symmetry Hamamelis spp. [Witchhazel]
• Anthers open by flaps in Hamamelis H. vernalis Sarg. [Spring or Vernal Witchhazel]
• Staminodes present in Hamamelis H. virginiana L. [American Witchhazel]
• Syncarpous gynoecium made up of 2 Hamamelis × intermedia ‘Arnold Promise’
connate carpels
[H. mollis × H. japonica]
• Inferior, half-inferior or superior ovary
Liquidambar styraciflua L. [Sweet Gum]
• 2-locular ovary with axile, pendulous
ovules Parrotia persica (DC.) C.A. Mey. [Persian Ironwood]

INFLORESCENCE TYPE(S) NOTE: Most species located in the North Temperate Zone

• Axillary clusters, spikes or heads

FRUIT TYPE(S)
• Woody loculicidal capsules
• 1 pendulous seed per locule
• Seed ejected in Hamamelis

HABIT
• Shrubs or trees
• Some wood resinous

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Toothed or lobed margin
• Stipules present
• Stellate hairs often present
• Often colored in autumn

IB 335 Plant Families 13


MORACEAE
mulberry or fig family
♂ Ca4 Co0 A4 G0

Eudicots: Rosid Clade ♀ Ca4 Co0 A0 G②

FLORAL CHARACTERISTICS EXAMPLES


• Tiny imperfect flowers Artocarpus spp. [Breadfruit]
• Plants monoecious or dioecious A. altilis (Parkinson) Fosberg [Breadfruit]
• Syncarpous gynoecium of 2 connate A. heterophyllus Lam. [Jackfruit]
carpels Broussonetia papyrifera (L.) L’Hér. ex Vent. [Paper Mulberry]
• Superior or inferior ovary Dorstenia spp. [Dorstenia]
• 1-locular ovary with 1 pendulous ovule Ficus spp. [Fig]
F. aurea Nutt. [Florida Strangler Fig]
INFLORESCENCE TYPE(S) F. benghalensis L. [Indian Banyan]
F. benjamina L. [Weeping Fig, Shopping Mall Fig]
• Flowers in heads, catkins or a SYCONIUM
F. carica L. [Edible Fig]
F. elastica Roxb. ex Hornem. [Indian Rubber Plant]
FRUIT TYPE(S) F. lyrata Warb. [Fiddleleaf Fig]
F. religiosa L. [Bo-Tree]
• Achenes or small drupes
Maclura pomifera (Raf.) C.K. Schneid. [Osage Orange]
• Sometimes multiple fruits with accessory
tissue [i.e. SYCONIUM] Morus spp. [Mulberry]

• Can be very large M. alba L. [White Mulberry, ‘Silkworm’ Mulberry]


M. rubra L. [Red Mulberry]

HABIT
• Trees and shrubs, sometimes herbs
• Sap milky

LEAF CHARACTERISTICS
• Simple leaves
• Alternate or opposite arrangement
• Entire, toothed or lobed margin

RELATED FAMILY: CANNABACEAE


Cannabis sativa L. [Marijuana]
Humulus lupulus L. [Common Hop]

14 Integrative Biology 335: Plant Systematics


FAGACEAE ♂ Ca 4-7 Co0 A4-∞ G0
beech or oak family
♀ Ca 4-7 Co0 A0
Eudicots: Rosid Clade
G③

FLORAL CHARACTERISTICS EXAMPLES


• Tiny imperfect flowers Castanea spp. [Chestnut]
• Plants usually monoecious C. dentata (Marsh.) Borkh. [American Chestnut]
• Actinomorphic symmetry C. mollissima Blume [Chinese Chestnut]
• Syncarpous gynoecium made up of 3 Fagus spp. [Beech]
connate carpels F. grandifolia Ehrh. [American Beech]
• Inferior ovary F. sylvatica L. [European Beech]
• 3-locular ovary with 2 pendulous ovules F. sylvatica ‘Atropunicea’ [Purple Beech]
in each locule [only 1 ovule per ovary
becoming a seed] F. sylvatica ‘Roseo-marginata’ [Tricolor Beech]
F. sylvatica ‘Pendula’
Quercus spp. [Oaks] - 2 subgenera in Illinois
INFLORESCENCE TYPE(S) Subgenera Erythrobalanus [Black/Red Oaks]
• Staminate flowers in catkins - Acorns mature in 2 years
• Carpellate flowers in few-flowered clusters - Leaves with bristle-tipped veins
enclosed by an INVOLUCRE Q. palustris Münchh. [Pin Oak]
Q. rubra L. [Northern Red Oak]
Q. velutina Lam. [Black Oak]
FRUIT TYPE(S)
Subgenera Quercus (Leucobalanus) [White Oaks]
• Nuts: 1-seeded and subtended or - Acorns mature in 1 year
surrounded by an INVOLUCRE [cupule]
- Leaves without bristle-tipped veins
Q. alba L. [White Oak] - State Tree of Illinois
HABIT Q. macrocarpa Michx. [Bur Oak]

• Trees or sometimes shrubs Other Oaks:

• Deciduous or evergreen [farther south] Q. robur L. [English Oak]


Q. suber L. [Cork Oak]
Q. virginiana Mill. [Live Oak]
LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Toothed or entire margin, or pinnately
lobed
• With stipules

IB 335 Plant Families 15


BETULACEAE ♂ Ca 0-6 Co0 A2-18 G0
birch family
♀ Ca 0-6 Co0 A0
Eudicots: Rosid Clade
G②

FLORAL CHARACTERISTICS EXAMPLES


• Tiny imperfect flowers Alnus spp. [Alders]
• Plants monoecious A. cordata (Loisel.) Duby [Italian Alder]
• Actinomorphic symmetry A. glutinosa (L.) Gaertn. [Common or Black Alder]
• Syncarpous gynoecium made up of 2 Betula spp. [Birches]
connate carpels B. nigra L. [River Birch]
• Inferior ovary [usually] *hard to see* B. papyrifera Marsh. [Paper or Canoe Birch]
• 2-locular ovary below but 1-locular above B. pendula Roth [European White Birch]
with 1 pendulous ovule per locule near top
of partition Carpinus spp. [Hornbeams]
C. betulus L. [European Hornbeam]
C. caroliniana Walter [American Hornbeam]
INFLORESCENCE TYPE(S) Corylus spp. [Hazelnuts, Filberts]
• Staminate flowers in 2-3 flowered cymules C. americana Walter [American Hazelnut or Filbert]
subtended by bracts and clustered into C. avellana L. [European Filbert]
catkins
C. avellana ‘Contorta’ [Harry Lauder’s Walking Stick]
• Carpellate flowers in 1-3 flowered cymules
subtended by bracts and clustered into Ostrya spp. [Hophornbeams]
catkins [these sometimes cone-like] O. virginiana (Mill.) K. Koch [American Hophornbeam]

FRUIT TYPE(S)
• Nuts and samaras that are 1-seeded

HABIT
• Trees and shrubs
• Mostly of cooler parts of the Northern
Hemisphere

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Toothed or doubly serrate margin
• Often with oblique leaf base
• Stipules present

16 Integrative Biology 335: Plant Systematics


CUCURBITACEAE ♂ Ca5 Co⑤ A⑤ G0
cucumber family
♀ Ca5 Co⑤ A0
Eudicots: Rosid Clade
G③

FLORAL CHARACTERISTICS EXAMPLES


• Imperfect flowers Citrullus lanatus (Thunb.) Matsum. & Nakai [Watermelon]
• Plants monoecious or dioecious Cucumis spp. [Melons]
• Actinomorphic symmetry C. melo L. [Cantelope]
• Petals connate and often yellow C. sativus L. [Garden Cucumber]
• Stamens [both filaments and anthers] often Cucurbita spp. [e.g. Pumpkin, Squash]
connate Ecballium elaterium (L.) A. Rich. [Squirting Cucumber]
• Anthers straight to folded or bent Echinocystis lobata (Michx.) Torr. & A. Gray [Wild Cucumber]
• Syncarpous gynoecium made up of 3 Lagenaria spp. [White-flowered Gourds]
connate carpels
Luffa aegyptiaca Mill. [Sponge Gourd]
• Inferior ovary
Marah macrocarpus (Greene) Greene [Cucamonga Manroot]
• 1-locular ovary with many parietal ovules
• Hypanthium sometimes present

INFLORESCENCE TYPE(S)
• Flowers solitary or in axillary cymes,
panicles or racemes

FRUIT TYPE(S)
• PEPOS, also berries or capsules

HABIT
• Herbaceous vines with tendrils
• Produce triterpenoid cucurbitacins

LEAF CHARACTERISTICS
• Alternate arrangement
• Palmately lobed or divided
• No stipules

IB 335 Plant Families 17


VIOLACEAE
violet family ♂ CaZ5 CoZ5 A5 G③

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• 2 types of flowers: CHASMOGAMOUS and Viola spp. [Violets]
CLEISTOGAMOUS V. pedata L. [Birdfoot Violet]
• Perfect flowers V. sororia Willd. [Common Blue Violet]
• Zygomorphic symmetry V. striata Aiton [Striped Cream Violet]
• 1 petal with SPUR V. tricolor L. [Pansy]
• 2 lower anthers, each with a nectary, Hybanthus concolor (T.F. Forst.) Spreng. [Eastern Greenviolet]
protrude into SPUR
• Syncarpous gynoecium made up of 3
connate carpels
• Superior ovary
• Ovary has many parietal ovules

INFLORESCENCE TYPE(S)
• Flowers solitary and scapose or in
racemes, panicles or cymes

FRUIT TYPE(S)
• Loculicidal capsules [explosively dehiscent
in Viola]
• Also berries or nuts

HABIT
• Annual or perennial herbs
• Shrubs and trees in the tropics

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Often toothed margin, sometimes lobed or
dissected
• Stipules present

18 Integrative Biology 335: Plant Systematics


SALICACEAE
♂ Ca0-x Co0 A2-∞ G0
willow family

Eudicots: Rosid Clade ♀ Ca 0-x Co0 A0 G 2-4

FLORAL CHARACTERISTICS EXAMPLES


• Simple imperfect flowers Salix spp. [Willows]
• Plants dioecious S. amygdaloides Andersson [Peachleaf Willow]
• Flowers often subtended by a bract S. babylonica L. [Weeping Willow]
• Calyx reduced to a disk in Populus or S. humilis Marsh. [Prairie Willow]
nectar glands in Salix Populus spp. [Poplars]
• Syncarpous gynoecium made up of 2-4 P. alba L. [White Poplar]
connate carpels
P. deltoides Bartram ex Marsh. [Eastern Cottonwood]
• Superior ovary
P. tremuloides Michx. [Quaking Aspen]
• 1-locular ovary with many parietal ovules
• Populus wind pollinated and Salix bee
pollinated

INFLORESCENCE TYPE(S)
• Catkins

FRUIT TYPE(S)
• Capsules
• Seeds hairy [comose]

HABIT
• Trees and shrubs
• Salicin [aspirin or acetylsalicylic acid]
originally from Salix

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Entire or toothed margin
• Stipules present

IB 335 Plant Families 19


FABACEAE (LEGUMINOSAE)
legume family ♂ Ca5 Co5 A10-∞ G1

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS HABIT


• Usually perfect flowers • Most legumes have root nodules containing nitrogen-fixing
• Monocarpous gynoecium bacteria

• Superior ovary • Enormous economic importance, second only to Poaceae


(Grass Family) for food
• Ovary has 2 to many marginal ovules that
occur in alternating rows on either side of
the carpel opening (suture)
LEAF CHARACTERISTICS
• Leaves alternate, often pinnately compound
FRUIT TYPE(S) • Less often leaves palmately compound or simple
• LEGUME: basically a follicle that opens
along two sutures rather than one (Note:
The fruit is a “legume” by definition, THREE SUBFAMILIES
but there is enormous variation in the
• MIMOSOIDEAE (Mimosa Subfamily)
morphology of fruits that are produced
from the single carpel) • CAESALPINIOIDEAE (Caesalpinia Subfamily)
• Seeds lack endosperm • FABOIDEAE (Bean or Pea Subfamily)

MIMOSOIDEAE CAESALPINIOIDEAE FABOIDEAE

Floral
Actinomorphic Zygomorphic Zygomorphic
Symmetry

1 BANNER petal outermost, 2


Corolla Petals connate and often forming BANNER petal internal to lateral
distinct lateral WING petals, and 2
Arrangement a tube petals
petals fused to form the KEEL

10 or more stamens 10 (or fewer) stamens 10 stamens either...


- DIADELPHOUS (usually)
Androecium Pollen in POLLINIA (aggregates of Anthers often opening by pores - MONADELPHOUS
pollen) - Distinct

Seed
U-shaped PLEUROGRAM O-shaped PLEUROGRAM No PLEUROGRAM
PLEUROGRAM

20 Integrative Biology 335: Plant Systematics


MIMOSOIDEAE
mimosa subfamily ♂ Ca⑤ Co⑤ A 10-∞ G1

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Acacia spp. [A very BIG genus]
• Actinomorphic symmetry Albizia julibrissin Durazz. [Silktree]
• Stamens usually 10 or more and pollen in Calliandra spp. [Powderpuff Tree]
pollinia Desmanthus illinoensis (Michx.) MacMill. [Illinois Bundleflower]
• Calyx, Corolla and filaments of Androecium Mimosa pudica L. [Sensitive Plant]
all connate forming tubes, but Ca, Co and
A free from one another
• Thus, flower has a “Tube in a Tube in a
Tube” structure

INFLORESCENCE TYPE(S)
• Capitate clusters, spikes or racemes

FRUIT TYPE(S)
• Typically a LEGUME [dry and dehiscing
along both sutures]
• Seeds without endosperm, often with a
U-shaped groove [PLEUROGRAM] or pits

HABIT
• Mostly tropical and subtropical trees
• A few temperate shrubs and herbs
• Usually with root nodules

LEAF CHARACTERISTICS
• Bipinnately compound leaves
• Alternate arrangement with a PULVINUS
• Some show movement
• Stipules present
• Beltian bodies in some species

IB 335 Plant Families 21


CAESALPINIOIDEAE
caesalpinia subfamily ♂ Ca5 CoZ5 A10 G1

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Bauhinia spp. [Orchid Tree]
• Zygomorphic symmetry Caesalpinia pulcherrima (L.) Sw. [Pride of Barbados]
• BANNER (standard) petal internal to 2 Ceratonia siliqua L. [Carob, St. John’s Bread]
distinct lateral (WING) petals and 2 distinct Cercis canadensis L. [Redbud]
petals forming the KEEL
Chamaecrista fasciculata (Michx.) Greene [Partridge Pea]
• Calyx, Corolla and Androecium usually
distinct Dalbergia spp. [Rosewood]

• Hypanthium present [too small to see] Delonix regia (Bojer ex Hook.) Raf. [Royal Poinciana]

• Stamens usually 10 [or fewer] and anthers Gleditsia triacanthos L. [Honey Locust]
often opening by pores Gymnocladus dioicus (L.) K. Koch [Kentucky Coffee tree]
Tamarindus indica L. [Tamarind]

INFLORESCENCE TYPE(S)
• Mostly racemes or spikes

FRUIT TYPE(S)
• LEGUMES [dry and dehiscing along both
sutures or breaking into LOMENTS]
• Seeds without endosperm, often with an
O-shaped groove [PLEUROGRAM]

HABIT
• Tropical and subtropical trees and shrubs
• A few temperate trees and herbs
• Usually without root nodules [only present
in 1/3 of species]

LEAF CHARACTERISTICS
• Pinnately or bipinnately compound leaves,
sometimes appearing simple
• Alternate arrangement with a PULVINUS
• Stipules present

22 Integrative Biology 335: Plant Systematics


FABOIDEAE
bean or pea subfamily ♂ Ca 5 CoZ 2
♀ +3
A 9 + 1 or 10 G1
Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Amorpha canescens Pursh [Lead Plant]
• Zygomorphic symmetry Arachis hypogaea L. [Peanut]
• 1 BANNER (standard) petal outermost, 2 Baptisia spp. [False Indigo]
distinct lateral WING petals and 2 petals Cladrastis kentukea (Dum. Cours.) Rudd [Yellowwood]
fused to form the KEEL
Coronilla varia (L.) Lassen [Crownvetch]
• Hypanthium present [too small to see]
Dalea spp. [Prairie Clover]
• 10 stamens usually DIADELPHOUS [9
connate filaments and 1 distinct filament]; D. purpurea Vent. var. purpurea [Purple Prairie Clover]
sometimes stamens MONADELPHOUS or all D. candida Michx. [White Prairie Clover]
distinct D. foliosa (A. Gray) Barneby [Leafy Prairie Clover]
Desmodium illinoense A. Gray [Tick Trefoil]
INFLORESCENCE TYPE(S) Erythrina herbacea L. [Redcardinal]
Glycine max (L.) Merr. [Soybean]
• Racemes, spikes or heads
Glycyrrhiza glabra L. [Cultivated Licorice]
Lathyrus spp. [Sweet Peas]
FRUIT TYPE(S) Lens culinaris Medik. [Lentil]
Lupinus spp. [Lupines]
• Diverse LEGUMES
Medicago spp. [Alfalfa]
• Seeds without endosperm and
PLEUROGRAM Phaseolus vulgaris L. [Common Bean]
Pisum sativum L. [Garden Pea]
Pueraria lobata (Willd.) Ohwi [Kudzu-vine]
HABIT Robinia pseudoacacia L. [Black Locust]
• Mostly herbs, some trees and shrubs Trifolium spp. [Clover]
• Temperate, subtropical and tropical
• Usually with root nodules

LEAF CHARACTERISTICS
• Usually pinnately compound, sometimes
palmately compound; rarely simple
• Alternate arrangement
• Stipules present

IB 335 Plant Families 23


ONAGRACEAE
evening primrose family ♂ Ca4 Co4 A8

Eudicots: Rosid Clade G④

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Clarkia spp. [Clarkia]
• Actinomorphic symmetry Epilobium spp. [Willowherb]
• Flowers 4-merous with hypanthium Fuchsia spp. [Fuchsia]
• Pollen sometimes with VISCIN THREADS Ludwigia spp. [False Loosestrife]
• Syncarpous gynoecium made up of 4 Oenothera biennis L. [Common Evening Primrose]
connate carpels
• Inferior ovary
• Often numerous axile or parietal ovules

INFLORESCENCE TYPE(S)
• Flowers solitary and axillary or in racemes
or spikes

FRUIT TYPE(S)
• Capsule or berry

HABIT
• Herbs or shrubs
• Sometimes aquatic

LEAF CHARACTERISTICS
• Simple leaves
• Alternate, opposite or whorled
arrangement
• No stipules

24 Integrative Biology 335: Plant Systematics


EUPHORBIACEAE
♂ Ca0 Co0 A1 G0
spurge family

Eudicots: Rosid Clade ♀ Ca 0 Co0 A0 G③

FLORAL CHARACTERISTICS EXAMPLES


• Imperfect flowers Acalypha spp. [Chenille plant, Copperleaf]
• Plants monoecious (usually) or dioecious Chamaesyce spp. [Creeping spurges]
• Actinomorphic symmetry Codiaeum variegatum (L.) A. Juss. [Garden Croton]
• Syncarpous gynoecium made up of 3 Croton spp. [Croton]
connate carpels Euphorbia spp. [BIG genus, often split into several smaller
• Superior ovary genera]
• 3-locular ovary with 1 apical ovule per E. corollata L. [Flowering Spurge]
locule E. fulgens Karw. ex Klotzsch [Scarlet-Plume]
E. marginata Pursh [Snow on the Mountain]
INFLORESCENCE TYPE(S) E. milii Des Moul. [Crown of Thorns]
E. pulcherrima Willd. ex Klotzsch [Poinsettia]
• A CYATHIUM with staminate flowers (consist
of only 1 stamen), carpellate flowers E. piscatoria Ait. [Figueira do inferno = Fig of Hell]
(consist of only a syncarpous gynoecium), Hevea brasiliensis (Willd. ex A. Juss.) Müll. Arg. [Rubber Tree]
nectar glands and petal-like appendages Jatropa podagrica Hook. [Gout plant]
Manihot esculenta Crantz [Cassava, Tapioca]
Ricinus communis L. [Casterbean]
FRUIT TYPE(S)
Triadica sebifera (L.) Small [Chinese tallow, Popcorn Tree]
• SCHIZOCARPS splitting into MERICARPS Vernicia fordii (Hemsl.) Airy-Shaw [Tungoil Tree]

HABIT
• Extremely variable - annual or perennial
herbs, shrubs or, in the tropics, large trees
• Often SUCCULENT and resembling cacti,
especially in southern Africa
• Often with milky sap
• Often poisonous

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement

IB 335 Plant Families 25


ROSACEAE
rose family

♀ Ca5 Co5 A∞ G ∞ or G1 or G 2-5
Eudicots: Rosid Clade

FLORAL CHARACTERISTICS HABIT


• Perfect, 5-merous flat flowers • Perennial herbs [rarely annuals], shrubs and trees
• Actinomorphic symmetry
• Hypanthium present, but variable
LEAF CHARACTERISTICS
• Sepals and petals distinct
• Petals round with wavy margins and never • Simple or compound (pinnate or palmate) leaves
really brilliant red or blue in wild • Alternate arrangement
• Stamens whorled • Usually stipules present
• Superior ovary [except in Subfamily
Maloideae]
FOUR SUBFAMILIES
• SPIRAEOIDEAE (Spirea Subfamily)
INFLORESCENCE TYPE(S)
• ROSOIDEAE (Rose Subfamily)
• Various • AMYGDALOIDEAE (Peach Subfamily)
• MALOIDEAE (Apple Subfamily)

FRUIT TYPE(S)
• Follicles, achenes, drupes or pomes (not
capsules or berries)
• Seeds usually without endosperm

SPIRAEOIDEAE ROSOIDEAE AMYGDALOIDEAE MALOIDEAE

Carpel Number 2-5 carpels that are


Few (5) distinct carpels Many distinct carpels 1 carpel
and Fusion fused to hypanthium

Ovary Position Superior ovary Superior ovary Superior ovary Inferior ovary

Achenes (in a HIP) or


Fruit Type Follicle Drupe POME
drupelets

Base Chromosome
9 7 8 17
Number

26 Integrative Biology 335: Plant Systematics


SPIRAEOIDEAE
spirea subfamily ♂
♀ Ca5 Co5 A10-∞ G 2-5
Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Aruncus dioicus (Walter) Fernald [Goat’s Beard]
• Actinomorphic symmetry Exochorda racemosa (Lindl.) Rehder [Pearlbush]
• Apocarpous gynoecium made up of 2-5 Physocarpus spp. [Ninebark]
distinct carpels Sorbaria sorbifolia (L.) A. Braun [False Spiraea]
• Superior ovary Spiraea spp. [Spirea, Meadowsweet, Hardhack, Steeple-bush]
• Marginal ovules Neillia sinensis Oliv. [Chinese Neillia]
• Base Chromosome Number = 9

FRUIT TYPE(S)
• Follicle [can have up to 5 follicles from one
flower]

HABIT
• Mostly shrubs, some herbs

LEAF CHARACTERISTICS
• Most have simple leaves, but Aruncus and
Sorbaria have pinnately compound leaves
• Alternate arrangement

IB 335 Plant Families 27


ROSOIDEAE
rose subfamily ♂
♀ Ca5 Co5 A10-∞ G ∞
Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Duchesnea indica (Andrews) Focke [Indian Strawberry]
• Actinomorphic symmetry Filipendula rubra (Hill) B.L. Rob. [Queen-of-the-Prairie]
• Apocarpous gynoecium made up of many Fragaria spp. [Strawberry]
distinct carpels Geum triflorum Pursh [Prairie Smoke]
• Superior ovary Kerria japonica (L.) DC. [Japanese Rose]
• Marginal ovules Potentilla spp. [Cinquefoil]
• Base Chromosome Number = 7 Rhodotypos scandens (Thunb.) Makino [Jetbead]
Rosa spp. [Rose]
FRUIT TYPE(S) Rubus spp. [Blackberry, Raspberry]
Sanguisorba spp. [Burnet]
• Achenes [in HIPS in Rosa]
Waldsteinia spp. [Barren Strawberry]
• Aggregate fruit of drupelets in Rubus

HABIT
• Shrubs and perennial herbs, rarely annuals

28 Integrative Biology 335: Plant Systematics


AMYGDALOIDEAE
peach subfamily ♂
♀ Ca5 Co5 A5-15 G1
Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Prinsepia spp. [Prinsepia]
• Actinomorphic symmetry Prunus spp. [Cherry, Peach, Plum, Apricot, Almond, Nectarine]
• Monocarpous gynoecium P. avium (L.) L. (Sweet Cherry)
• Superior ovary P. dulcis (Mill.) D.A. Webb (Sweet Almond)
• Marginal ovules P. persica (L.) Batsch (Peach)
• Base Chromosome Number = 8 Maddenia spp. [Maddenia]

FRUIT TYPE(S)
• Drupe

HABIT
• Trees and shrubs [NO herbs]
• Bark often beautiful with horizontal
LENTICELS

IB 335 Plant Families 29


MALOIDEAE
apple subfamily ♂
♀ Ca5 Co5 A10-∞

Eudicots: Rosid Clade G 2-5

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Amelanchier spp. [Shadbush]
• Actinomorphic symmetry Aronia spp. [Chokeberry]
• Syncarpous gynoecium made up of 2-5 Chaenomeles spp. [Japanese Quince]
[somewhat] connate carpels Cotoneaster spp. [Cotoneaster]
• Half to completely inferior ovary Crataegus spp. [Hawthorn]
• Ovary partly to completely adnate to Cydonia spp. [Quince]
hypanthium
Malus spp. [i.e. Apple, Crabapple]
• Base Chromosome Number = 17
Pyracantha spp. [Firethorn]
Pyrus spp. [Pear]
FRUIT TYPE(S) Sorbus spp. [Mountain Ash]

• POME with a core or with small stones

HABIT
• Trees and shrubs

30 Integrative Biology 335: Plant Systematics


BRASSICACEAE (CRUCIFERAE)
mustard family
♂ Ca4 Co4 A4+2 G②

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Alliaria petiolata (M. Bieb.) Cavara & Grande [Garlic Mustard]
• Actinomorphic symmetry Armoracia rusticana G. Gaertn., B. Mey. & Scherb.
• 4-merous flowers [traditional family name [Horseradish]
is Cruciferae meaning “cross”] Barbarea vulgaris W.T. Aiton [Garden Yellowrocket]
• TETRADYNAMOUS stamens: 4 long and 2 Brassica spp. [Mustards]
short stamens B. campestris L. [e.g. Turnips, Chinese cabbage]
• Syncarpous gynoecium made up of 2 B. juncea (L.) Czern. [India mustard]
connate carpels
B. nigra (L.) W.D.J. Koch [Black mustard]
• Superior ovary
B. napus L. [Rape seed]
• 2-locular ovary with many parietal ovules
B. oleracea L. [Cabbage]
B. oleracea var. botrytis L. [Broccoli]
INFLORESCENCE TYPE(S) B. oleracea var. gemmifera DC. [Brussels sprouts]

• Bractless racemes Capsella bursa-pastoris (L.) Medik. [Shepherd’s purse]


Cardamine concatenata (Michx.) Sw. [Cutleaf Toothwort]
Hesperis matronalis L. [Dames Rocket]
FRUIT TYPE(S) Iberis spp. [Candytuft]

• SILIQUES or SILICLES with a REPLUM Matthiola spp. [Stocks]


[siliques or silicles may be flattened Lepidium spp. [Pepperweeds]
perpendicular or parallel to the replum] Lesquerella ludoviciana (Nutt.) S. Watson [Bladderpod]
Lunaria annua L. [Money plant, Honesty]
HABIT Nasturtium officinale W.T. Aiton [Watercress]
Raphanus sativus L. [Cultivated Radish]
• Annual, biennial or perennial herbs, very
rarely shrubs
• Produce mustard oil glucosides

LEAF CHARACTERISTICS
• Simple or compound leaves
• Alternate arrangement
• May be pinnately lobed
• No stipules

IB 335 Plant Families 31


MALVACEAE
mallow family A ∞
♂ Ca⑤ Co5 G 5-∞

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Abelmoschus esculentus (L.) Moench [Okra]
• Actinomorphic symmetry Abutilon theophrasti Medik. [Velvetleaf]
• Sepals often subtended by bracts Alcea rosea L. [Hollyhock]
[EPICALYX] Althaea officinalis L. [Common Marshmallow]
• Petals distinct but adnate to the staminal Callirhoë spp. [Poppy Mallow]
tube
Gossypium spp. [Cotton]
• MONADELPHOUS stamens - the filaments of
the stamens are connate forming a tube Hibiscus spp. [Rosemallow]
around the gynoecium H. syriacus L. [Rose-of-Sharon]
• Anthers unilocular and crescent-shaped Iliamna remota Greene [Kankakee Mallow]
• Syncarpous gynoecium made up of many Napaea dioica L. [Glademallow]
connate carpels
• Superior ovary enclosed by staminal tube
• Many locules with many axile ovules

INFLORESCENCE TYPE(S)
• Flowers solitary or in cymes

FRUIT TYPE(S)
• Schizocarps or capsules
• Berries

HABIT
• Herbs and shrubs, rarely trees
• Stellate hairs common
• Often mucilaginous sap

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Often toothed and palmately lobed and
veined

32 Integrative Biology 335: Plant Systematics


ACERACEAE ♂
maple family ♂
♀ or ♀ Ca5(4) Co5(4) A8,10,5 * G② †
Eudicots: Rosid Clade Rudimentary in ♀ flowers (*) and ♂ flowers (†)

FLORAL CHARACTERISTICS EXAMPLES


• Imperfect (usually) or perfect flowers Acer spp. [Maples]
• If imperfect, plants monoecious or A. ginnala Maxim. [Amur maple]
dioecious A. griseum (Franch.) Pax [Paperbark maple]
• Actinomorphic symmetry A. negundo L. [Box-elder]
• Syncarpous gynoecium made up of 2 A. nigrum Michx. f. [Black maple]
connate carpels
A. palmatum Thunb. [Japanese maple]
• Superior winged ovary
A. pensylvanicum L. [Striped maple, Moosewood]
• 2-locular ovary with 2 axile ovules per
locule A. platanoides L. [Norway maple]

• Wind or insect pollinated [latter with nectar A. pseudoplatanus L. [Sycamore maple]


disk] A. rubrum L. [Red maple]
A. saccharinum L. [Silver maple]
A. saccharum Marsh. [Sugar maple]
INFLORESCENCE TYPE(S)
[40 gallons of sap = 1 gallon of syrup]
• Various

FRUIT TYPE(S)
• SAMAROID SCHIZOCARPS [Schizocarps
break into 2 samaras]

HABIT
• Deciduous trees and some shrubs

LEAF CHARACTERISTICS
• Simple palmately lobed [divided] leaves or
compound leaves [pinnately compound in
Acer negundo]
• Opposite arrangement
• No stipules

IB 335 Plant Families 33


ERICACEAE ♂ Ca5 Co⑤ A10 G 2-10

heather family
OR
♂ Ca5 Co⑤ A10

Eudicots: Asterid Clade
G⑤

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Arbutus spp. [Madrone]
• Actinomorphic symmetry [zygomorphic in Arctostaphylos spp. [Bearberry, Manzanita]
Rhododendron] Calluna and Erica spp. [Heathers]
• Perianth often URCEOLATE or Calluna vulgaris (L.) Hull [Heather]
CAMPANULATE
Erica tetralix L. [Crossleaf Heath]
• Anthers often APPENDAGED and opening
by pores Kalmia angustifolia L. [Sheep Laurel]

• Pollen often in tetrads Oxydendrum arboreum (L.) DC. [Sourwood]

• Syncarpous gynoecium; style hollow Pieris spp. [Fetterbush]

• Superior or inferior ovary Rhododendron spp. [e.g. Rhododendron, Azaleas]

• Ovary has as many locules as carpels with Rhododendrons are usually everygreen while Azaleas are
many axile ovules mostly deciduous [exceptions to both do occur].
Vaccinium spp. [e.g. Blueberries, Cranberries]

INFLORESCENCE TYPE(S)
• Various Pyrola spp. [Wintergreen]
Chimaphila spp. [Pipsissewa]
These two genera are often placed in a separate family,
FRUIT TYPE(S) Pyrolaceae.

• Capsules, berries or drupes


Monotropa spp. [Indian Pipe]
This genus and related genera, which are saprophytic, are
HABIT often placed in a separate family, Monotropaceae.

• Mainly small to large, evergreen or


deciduous shrubs
• Prefers acidic soil

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Mostly entire margin
• Often thick, leathery and evergreen
• No stipules

34 Integrative Biology 335: Plant Systematics


ASCLEPIADACEAE
milkweed family A⑤
♂ Ca⑤ Co⑤ G 2

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Asclepias spp. [Milkweeds]
• Actinomorphic symmetry A. curassavica L. [Bloodflower]
• Petals often with HORN and HOOD that A. incarnata L. [Swamp Milkweed]
make up the CORONA A. sullivantii Engelm. ex A. Gray [Sullivant’s Milkweed]
• Pollen in pollinia and pollinia of adjacent A. syriaca L. [Common Milkweed]
anther sacs connected by TRANSLATOR
A. tuberosa L. [Butterfly Milkweed]
ARMS with gland or CORPUSCULUM
A. verticillata L. [Whorled Milkweed]
• Androecium [filaments and anthers] and
stigma fused into a GYNOSTEGIUM Ceropegia sandersonii Decne. ex Hook. [Parachute plant]
• Syncarpous gynoecium made up of 2 Hoya carnosa (L. f.) R. Br. [Porcelainflower]
carpels that are distinct below Stapelia hirsuta L. [Carrion flower]
• Superior ovary Stephanotis floribunda Brongn. [Madagascar jasmine]
• Ovary with many marginal ovules

INFLORESCENCE TYPE(S)
• Umbels or cymes
• Flowers solitary in succulent species

FRUIT TYPE(S)
• Follicles
• Seeds with a COMA of hairs

HABIT
• Perennial herbs, vines, often trees and
shrubs in the tropics and succulents in
South Africa
• Milky sap

LEAF CHARACTERISTICS
• Simple leaves
• Opposite or whorled arrangement
• Entire margin
• No stipules

IB 335 Plant Families 35


SOLANACEAE
potato family A5
♂ Ca⑤ Co⑤ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Atropa bella-donna L. [Atropine]
• Actinomorphic symmetry [or weakly Browallia spp. [Browallia]
zygomorphic] Capsicum spp. [Peppers]
• Corolla PLICATE Datura spp. [Jimsonweed]
• Epipetalous stamens [stamens adnate to Nierembergia spp. [Cupflower]
petal tube]
Nicotiana tabacum L. [Tobacco]
• Anthers often CONNIVENT
Petunia × hybrida [Petunia]
• Syncarpous gynoecium made up of 2
connate carpels Physalis spp. [Chinese lanterns]

• Superior ovary P. ixocarpa Brot. [Tomatillo]

• 2-locular ovary with many axile ovules Schizanthus spp. [Butterfly flower]
Solanum spp. [e.g. Nightshades, Potatoes, Eggplant]
S. lycopersicum L. [Garden Tomato]
INFLORESCENCE TYPE(S) S. melongena L. [Eggplant]
• Flowers in cymes or solitary

FRUIT TYPE(S)
• Berries or capsules

HABIT
• Mostly herbaceous
• Some shrubs, woody vines and trees in
the tropics
• Some species poisonous, medicinal or
hallucinogenic

LEAF CHARACTERISTICS
• Simple leaves
• Alternate arrangement
• Entire or variously lobed to pinnate margin
• No stipules

36 Integrative Biology 335: Plant Systematics


OLEACEAE
olive family A2
♂ Ca④ Co④ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect or imperfect flowers Chionanthus spp. [Fringe tree]
• If imperfect flowers, plants dioecious Forsythia spp. [Forsythia]
• Actinomorphic symmetry Fraxinus spp. [Ash]
• 4-merous flowers Jasminum grandiflorum L. [Jasmine]
• 2 epipetalous stamens Ligustrum spp. [Privet]
• Syncarpous gynoecium made up of 2 Olea europaea L. [Olive]
connate carpels Syringa spp. [Lilac]
• Superior ovary
• 2-locular ovary with 2 axile ovules in each
locule [except 4-10 in Forsythia]

INFLORESCENCE TYPE(S)
• Racemes or panicles

FRUIT TYPE(S)
• Berry, capsule, drupe or samara

HABIT
• Trees and shrubs

LEAF CHARACTERISTICS
• Simple or pinnately compound leaves
• Opposite arrangement
• No stipules

IB 335 Plant Families 37


SCROPHULARIACEAE
figwort family A2 or 4 (5)
♂ CaZ⑤ CoZ⑤ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Antirrhinum spp. [Snapdragon]
• Zygomorphic symmetry [2-lipped] to nearly Calceolaria spp. [Slipper Flower]
actinomorphic Castilleja spp. [Indian Paintbrush]
• Sepals and petals each connate Chelone spp. [Turtlehead]
• Epipetalous stamens Collinsia verna Nutt. [Blue-eyed Mary]
• 2 or 4 DIDYNAMOUS stamens Digitalis purpurea L. [Purple Foxglove]
• 5th STAMINODE sometimes present Linaria spp. [Butter & Eggs, Toadflax]
• Syncarpous gynoecium made up of 2 Pedicularis spp. [Lousewort]
connate carpels
Penstemon spp. [Beards-tongue]
• Superior ovary
Striga spp. [Witchweed]
• 2-locular ovary with many axile ovules
Verbascum spp. [Mullein]
Veronicastrum virginicum (L.) Farw. [Culver’s Root]
INFLORESCENCE TYPE(S)
• Various cymes and racemes OROBANCHACEAE: parasitic, no chlorophyll
BIGNONIACEAE: mostly tropical trees and lianas

FRUIT TYPE(S)
• 2-valved capsules
• Seeds are tiny

HABIT
• Annual or perennial herbs, shrubs, trees or
woody vines
• Many photosynthetic root parasites

LEAF CHARACTERISTICS
• Simple to divided or compound leaves
• Alternate, opposite or whorled
arrangement
• No stipules

38 Integrative Biology 335: Plant Systematics


LAMIACEAE
mint family A4
♂ CaZ⑤ CoZ⑤ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Coleus spp. [Coleus]
• Zygomorphic symmetry Glechoma hederacea L. [Ground Ivy]
• Calyx and Corolla tubes each present Lamium spp. [e.g. Henbit, Dead nettle]
[Corolla bilabiate: 2-lobed upper, 3-lobed Lavendula spp. [Lavender]
lower]
Mentha spp. [Mints]
• Epipetalous stamens
M. arvensis L. [Menthol]
• 2 + 2 DIDYNAMOUS stamens
M. piperita L. [Peppermint]
• Syncarpous gynoecium made up of 2
connate carpels; deeply 4-lobed with M. spicata L. [Spearmint]
GYNOBASIC style Monarda fistulosa L. [Beebalm, Horsemint]
• Superior ovary Nepeta cataria L. [Catnip]
• 4-locular ovary with 1 basal/axile ovule per Ocimum basilicum L. [Basil]
locule Origanum majorana L. [Marjoram]
Origanum vulgare L. [Oregano]
INFLORESCENCE TYPE(S) Physostegia virginiana (L.) Benth. [False Dragonshead]
Rosmarinus officinalis L. [Rosemary]
• Axillary cymes or VERTICILS
Salvia officinalis L. [Kitchen Sage]
Satureja hortensis L. [Summer Savory]
FRUIT TYPE(S) Teucrium canadense L. [Canada Germander]
Thymus vulgaris L. [Garden Thyme]
• 4 nutlets
• Pericarp tightly surrounds each seed

HABIT
• Aromatic annual or perennial herbs
• Rarely trees, shrubs or woody vines
• Stems square in cross-section

LEAF CHARACTERISTICS
• Simple leaves
• Opposite or whorled arrangement
• No stipules
• Glandular trichomes with essential oils
[e.g. menthol]

IB 335 Plant Families 39


CAPRIFOLIACEAE
A 5
honeysuckle family ♂ Ca 5 Co⑤

Eudicots: Asterid Clade G 2-5

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Diervilla spp. [Bush Honeysuckle]
• Actinomorphic or zygomorphic symmetry Kolkwitzia amabilis Graebn. [Beautybush]
and sometimes 2-lipped Linnaea borealis L. [Twinflower]
• Epipetalous stamens Lonicera spp. [Honeysuckle]
• Syncarpous gynoecium made up of 2-5 Sambucus spp. [Elderberry]
connate carpels
Triosteum spp. [Horse Gentian]
• Inferior ovary
Viburnum spp. [e.g. Cranberrybush, Nannyberry]
• 2 to 5-locular ovary with 1 to few apical
ovules per locule
• Ovaries of 2 flowers often paired

INFLORESCENCE TYPE(S)
• Cymes, corymbs, panicles or in axillary
clusters

FRUIT TYPE(S)
• Berries, capsules or drupes

HABIT
• Shrubs, woody vines and small trees
• Rarely herbs

LEAF CHARACTERISTICS
• Simple or pinnately compound leaves
• Opposite arrangement
• No stipules

40 Integrative Biology 335: Plant Systematics


ASTERACEAE
A 5
sunflower family ♂ CaP Co⑤

Eudicots: Asterid Clade G②

FLORAL CHARACTERISTICS EXAMPLES: 3 TYPES OF CAPITULA [HEADS]


• Perfect or imperfect flowers
• If imperfect flowers, plants monoecious RADIATE CAPITULUM: Both DISK and RAY flowers
• Two kinds of flowers:
Aster spp. [Aster]
DISK: Actinomorphic, tubular corolla
Bidens spp. [Beggar-ticks]
RAY: Zygomorphic, ligulate corolla
Chrysanthemum spp. [Daisy]
• Calyx = PAPPUS; Bracts = CHAFF
Echinacea, Ratibida, Rudbeckia spp. [Coneflowers]
• Epipetalous stamens; stamen filaments
distinct but anthers connate into a tube Helianthus spp. [Sunflowers]
around style Senecio spp. [Ragworts]
• Syncarpous gynoecium made up of 2 Silphium spp. [Rosinweed]
connate carpels S. laciniatum L. [Compass Plant]
• Inferior ovary S. perfoliatum L. [Cup Plant]
• 1-locular ovary with 1 basal ovule S. terebinthinaceum Jacq. [Prairie Dock]
Solidago spp. [Goldenrods]
INFLORESCENCE TYPE(S) Tagetes spp. [Marigolds]
Zinnia spp. [Zinnia]
• Heads [CAPITULUM, CAPITULA] surrounded
by PHYLLARIES
• Heads arranged in secondary cymes or
racemes DISCOID CAPITULUM: DISK flowers only
Arctium spp. [Burdock]
FRUIT TYPE(S) Cirsium & Carduus spp. [Thistles]
Cynara scolymus L. [Globe Artichoke]
• Achenes
Eupatorium spp. [Joe-Pyeweed]
Liatris spp. [Blazing Stars]
HABIT Vernonia spp. [Ironweeds]

• Mostly annual, biennial or perennial herbs


• Shrubs or tree-like at high elevations in the
tropics and on some islands LIGULATE CAPITULUM: RAY flowers only
Cichorium spp. [e.g. Chicory, Endive]
LEAF CHARACTERISTICS Hieracium spp. [Hawkweed]
Krigia spp. [Dwarf Dandelion]
• Mostly simple leaves [often lobed or
divided], rarely compound Lactuca spp. [Lettuce]
• Alternate, opposite or whorled Taraxacum spp. [Dandelion]
arrangement Tragopogon spp. [Goat’s Beard]

IB 335 Plant Families 41


42 Integrative Biology 335: Plant Systematics
APIACEAE (UMBELLIFERAE)

carrot family ♂
♀ or ♀ Ca5 Co5 A5
Eudicots: Asterid Clade G②

FLORAL CHARACTERISTICS EXAMPLES


• Imperfect (usually) or small perfect flowers Aegopodium podagraria L. [Bishop’s Goutweed]
• If imperfect, plants monoecious or rarely Anethum graveolens L. [Dill]
dioecious Apium graveolens L. [Celery]
• Actinomorphic symmetry with a trend to Astrantia major L. [Greater Masterwort]
zygomorphy
Carum carvi L. [Caraway]
• Syncarpous gynoecium made up of 2
connate carpels; with a STYLOPODIUM Cicuta bulbifera L. [Bulblet-bearing Water Hemlock]

• Inferior ovary Conium maculatum L. [Poison Hemlock]

• 2-locular ovary with 1 pendulous ovule per Coriandrum sativum L. [Coriander, Cilantro]
locule Crithmum maritimum L. [Rock Samphire]
Cuminum cyminum L. [Cumin]
Daucus carota L. [Carrot, Queen Anne’s lace]
INFLORESCENCE TYPE(S)
Eryngium yuccifolium Michx. [Rattlesnake Master]
• Simple or compound umbels Foeniculum vulgare Mill. [Sweet Fennel]
Heracleum spp. [Cow Parsnip, Hogweed]
Hydrocotyle bonariensis Comm. ex Lam. [Pennywort]
FRUIT TYPE(S)
Monizia edulis Lowe [Tree carrot]
• Ribbed schizocarps = 2 mericarps that are Osmorhiza longistylis (Torr.) DC. [Sweet cicely]
connected by a CARPOPHORE
Pastinaca sativa L. [Wild Parsnip]
• Contain oil tubes
Petroselinum crispum (Mill.) Nyman ex A.W. Hill [Parsley]
Pimpinella anisum L. [Anise Burnet Saxifrage]
HABIT Trachymene spp.

• Annual, biennial or perennial herbs with Zizia aurea (L.) W.D.J. Koch [Golden Alexander]
hollow stems
• Aromatic and sometimes deadly poisonous

LEAF CHARACTERISTICS
• Most pinnately compound leaves
• Alternate arrangement
• Sheathing leaf base
• Have oil tubes

IB 335 Plant Families 43


ARACEAE
arum family ♂

♀ or ♀ Ca0 or 2-3 Co 0 or 2-3 A4-10 G 2 - 4
Monocot Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect or imperfect flowers Amorphophallus titanum (Becc.) Becc. ex Arcang [Titan Arum]
• If imperfect, plants mainly monoecious Anthurium spp. [Laceleaf]
• Actinomorphic symmetry Arisaema triphyllum (L.) Schott [Jack-in-the-pulpit]
• Flowers very reduced, often sunken into Caladium spp. [Caladium]
SPADIX Colocasia esculenta (L.) Schott [Taro root, Poi]
• Syncarpous gynoecium made up of 2 to 4 Dieffenbachia spp. [Dumb cane]
connate carpels
Monstera spp. [Monstera]
• Superior ovary
Philodendron spp. [Philodendron]
• 1 to many-locular ovary with 1 to many
ovules Spathiphyllum spp. [Spath or Peace Lilies]
Symplocarpus foetidus (L.) Salisb. ex Nutt. [Skunk cabbage]
Zantedeschia spp. [Calla lily]
INFLORESCENCE TYPE(S)
• A SPADIX subtended by a SPATHE
• Can be large [e.g. Amorphophalus]

FRUIT TYPE(S)
• Multiple, often colorful berries

HABIT
• Perennial herbs or woody vines
• Many species are shade tolerant

LEAF CHARACTERISTICS
• Simple to pinnately or palmately veined,
divided or compound
• Alternate arrangement
• Often with RAPHIDES of calcium oxalate,
poisonous

44 Integrative Biology 335: Plant Systematics


ARECACEAE (PALMAE)
palm family ♂

♀ or ♀ Ca3 Co3 A6 G 3 or ③
Monocot Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect or imperfect [small] flowers Cocos nucifera L. [Coconut Palm]
• If imperfect, plants monoecious or Corypha umbraculifera L. [Talipot Palm]
dioecious Elaeis guineensis Jacq. [African Oil Palm]
• Actinomorphic symmetry Lodoicea maldivica (J.F. Gmel.) Pers. [Double Coconut]
• Apocarpous gynoecium made up of 3 Phoenix dactylifera L. [Date Palm]
distinct carpels or Syncarpous gynoecium
made up of 3 connate carpels Roystonea elata (Bartram) F. Harper [Royal Palm]

• Superior ovary Sabal palmetto (Walter) Lodd. ex Schult. & Schult. f. [Palmetto]

• 1 to 3-locular ovary with 1 basal ovule per Serenoa repens (Bartram) Small [Saw Palmetto]
locule Washingtonia filifera (Linden ex André) H. Wendl. [California
Fan Palm]

INFLORESCENCE TYPE(S)
• Various spikes or panicles, sometimes
terminal and the plant monocarpic
• Can be gigantic (3,000,000 flowers and
250,000 fruits)

FRUIT TYPE(S)
• Drupe or berry
• 1-seeded
• Can be enormous in size [up to 18” long
and 40 lbs in weight]

HABIT
• Unbranched trees, shrubs or woody vines,
but without secondary growth

LEAF CHARACTERISTICS
• Often pinnately or palmately compound
leaves
• Alternate arrangement
• Sheathing leaf bases
• Sometime huge leaves [up to 65 ft long]

IB 335 Plant Families 45


ORCHIDACEAE
orchid family ♂ Ca3 CoZ 3 A1 or (2) (3)

G③
Monocot Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers TEMPERATE SPECIES
• Zygomorphic symmetry
Calopogon spp. [Grass Pink]
• RESUPINATE Corolla with 3 petals where
one is modified into a LIP [LABELLUM] Corallorhiza spp. [Coral-root]

• A and G adnate into a COLUMN Cypripedium spp. [Lady’s Slipper]

• Pollen in POLLINIA with a CAUDICLE and Goodyera spp. [Rattlesnake Plantain]


VISCIDIUM Habenaria spp. [Fringed Orchid]
• Part of stigma modified into ROSTELLUM Orchis spp. [Orchids]
• Syncarpous gynoecium made up of 3 Spiranthes spp. [Ladies’-tresses]
connate carpels
• Inferior [pedicel-like] ovary TROPICAL SPECIES
• 1-locular ovary with many parietal ovules
Angraecum spp. [Star-of-Bethlehem Orchid]
Cattleya spp. [Cattleya]
INFLORESCENCE TYPE(S) Cymbidium spp. [Cymbidium]
• Spikes, racemes, panicles or flowers Epidendrum spp. [Star Orchid]
solitary on a scape Oncidium spp. [Dancing-lady Orchid]
Paphiopedilum spp. [Venus’ slipper]
Phalaenopsis spp. [Moth orchid]
FRUIT TYPE(S)
Vanda spp. [Vanda]
• Loculicidal capsules Vanilla planifolia Jacks. [Vanilla ‘beans’]
• Countless seeds are microscopic and often
require a fungal symbiont to germinate

HABIT
• Perennial herbs
• Epiphytic or terrestrial, some saprophytic
• Adventitious roots with VELAMEN covering

LEAF CHARACTERISTICS
• Simple, entire [often rather fleshy] leaves
• Alternate or basal arrangement
• In epiphytic species the blade often
attached to a PSEUDOBULB

46 Integrative Biology 335: Plant Systematics


IB 335 Plant Families 47
IRIDACEAE
iris family ♂ Ca3 Co3 A3

G③
Monocot Clade

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Belamcanda chinensis (L.) DC. [Blackberry Lily]
• Actinomorphic or zygomorphic symmetry Crocus spp. [Crocus]
• Hypanthium often present C. angustifolius Weston [Cloth of Gold]
• Petaloid sepals and showy petals C. sativus L. [Saffron = 13,000 stigmas per ounce]
• Style petaloid in Iris spp. Freesia spp. [Freesia]
• 3 stamens in Androecium Gladiolus spp. [Gladiolus]
• Syncarpous gynoecium made up of 3 Iris spp. [for Greek Goddess of the Rainbow]
connate carpels Sisyrinchium spp. [Blue-eyed Grass]
• Inferior ovary Tigridia spp. [Peacock Flower]
• 3-locular ovary with many axile ovules per
locule

INFLORESCENCE TYPE(S)
• Cymes, umbels, panicles or flowers
solitary on a scape

FRUIT TYPE(S)
• Loculicidal capsules

HABIT
• Perennial herbs with bulbs, rhizomes or
corms

LEAF CHARACTERISTICS
• Simple leaves
• Alternate or basal arrangement
• Entire margin
• EQUITANT leaves [2 ranked and folded in
half lengthwise]
• Sheathing leaf bases

48 Integrative Biology 335: Plant Systematics


LILIACEAE ♂ Ca3 Co3 A6 G③

lily family
OR
♂ Ca3 Co3 A6

Monocot Clade
G③

FLORAL CHARACTERISTICS EXAMPLES


• Perfect flowers Allium spp. [e.g. Onion, Garlic, Leek]
• Actinomorphic symmetry Asparagus officinalis L. [Garden Asparagus]
• Sepals and petals similar [thus both called Clintonia borealis (Aiton) Raf. [Bluebead Lily]
TEPALS] except in Trillium spp. Convallaria majalis L. [European Lily of the Valley]
• TEPALS distinct or connate Erythronium spp. [e.g. Trout or Fawn Lily, Dog-tooth Violet]
• 6 stamens with often VERSATILE anthers Fritillaria imperialis L. [Crown Imperial]
• Syncarpous gynoecium made up of 3 Gloriosa spp. [Gloriosa Lily]
connate carpels
Hawthornia spp. [Star cactus]
• Superior or inferior ovary
Hyacinthus spp. [Hyacinth]
• 3-locular ovary with 1 to many axile ovules
per locule Kniphofia spp. [Redhot poker]
Lilium spp. [Lily]
Liriope spp. [Lilyturf]
INFLORESCENCE TYPE(S) Muscari botryoides (L.) Mill. [Common Grape Hyacinth]
• Umbels, panicles, racemes or flowers Ornithogalum umbellatum L. [Star-of-Bethlehem]
solitary on a scape Polygonatum spp. [Solomons seal]
Ruscus aculeatus L. [Butcher’s broom]
Smilacina spp. [False Solomon’s seal]
FRUIT TYPE(S)
Smilax spp. [Greenbrier]
• Capsules and berries Trillium spp. [Wake-robin]
Tulipa spp. [Tulip]
HABIT Uvularia spp. [Bellwort]
Yucca spp. [Yucca]
• Perennial herbs often with bulbs, rhizomes
or corms

LEAF CHARACTERISTICS
• Simple leaves
• Alternate or basal arrangement, rarely
whorled
• Entire margin
• Often sheathing leaf bases
• Leaves reduced to scales in Asparagus

IB 335 Plant Families 49


POACEAE
grass family ♂

♀ or ♀ Ca2 Co 0 A3 G 2-3
Monocot Clade

FLORAL CHARACTERISTICS LEAF CHARACTERISTICS [CONTINUED]


• Perfect or imperfect [reduced] flowers • Blades flat and sheathing at base with sheath open
• If imperfect, plants monoecious or • A LIGULE often present at junction of blade and sheath
dioecious
• Basic unit a FLORET [a flower subtended by
a PALEA and LEMMA] EXAMPLES
• Perianth represented by LODICULES
Andropogon gerardii Vitman [Big Bluestem]
• Anthers VERSATILE
Avena sativa L. [Common Oat]
• Syncarpous gynoecium made up of 2 or 3
connate carpels Bambusa spp. [Bamboo]
• Superior ovary Digitaria spp. [Crab grass]
• 1-locular ovary with 1 basal ovule Elymus spp. [Wild rye]
Festuca spp. [Fescue]
Hordeum vulgare L. [Common Barley]
INFLORESCENCE TYPE(S)
Miscanthus sinensis ‘Zebrinus’ [Zebra Grass]
• SPIKELETS grouped into panicles, racemes Oryza sativa L. [Rice]
or spikes
Panicum spp. [Panic grass, Switchgrass]
• Each SPIKELET has a RACHILLA as the
Poa spp. [Bluegrass]
central axis and two GLUMES at the base
Saccharum officinarum L. [Sugarcane]
Secale cereale L. [Cereal rye]
FRUIT TYPE(S) Sorghum bicolor (L.) Moench [Sorghum]
Sporobolus heterolepis (A. Gray) A. Gray [Dropseed]
• CARYOPSIS [an achene with the pericarp
fused to the seed coat] Triticum aestivum L. [Common wheat]
T. durum Desf. [Durum wheat]
Zea mays L. [Maize, corn]
HABIT
Zizania palustris L. [Northern Wild rice]
• Annual or perennial herbs
• Sometimes quite large [i.e. bamboo]
• Form bunches or with stolons or rhizomes
• Stems [CULMS] round in cross section with
solid nodes and hollow internodes

LEAF CHARACTERISTICS
• Simple, 2-ranked, entire or finely toothed,
linear leaves
• Alternate or basal arrangement

50 Integrative Biology 335: Plant Systematics


IB 335 Plant Families 51
52 Integrative Biology 335: Plant Systematics
LABORATORY 1
WOODY TWIG TERMINOLOGY

WOODY TWIG TERMINOLOGY LAB EXERCISE

The identification of woody plants in winter is quite different from the techniques used for the determination of the
same plants during the other seasons of the year. The features used are, of course, not usually those of the flowers
and fruits, or even of leaves, but rather the size and form of the plants and especially characteristics of the twigs. In
this Lab, you will gain experience in using a dichotomous key and learning woody twig vegetative terminology while
identifying woody plants in non-flowering condition. To complete this Lab Exercise, work through the following steps:

1. Read “Dichotomous Key Guidelines” and then identify all the twig specimens on display using the “Dichotomous
(Winter) Key for Woody Plants in Champaign-Urbana”.

Specimen A: ______________________________________________________________________________

Specimen B: ______________________________________________________________________________

Specimen C: ______________________________________________________________________________

Specimen D: ______________________________________________________________________________

Specimen E: ______________________________________________________________________________

Specimen F: ______________________________________________________________________________

Specimen G: ______________________________________________________________________________

Specimen H: ______________________________________________________________________________

Specimen I: ______________________________________________________________________________

Specimen J: ______________________________________________________________________________

Specimen K: ______________________________________________________________________________

Specimen L: ______________________________________________________________________________

Specimen M: ______________________________________________________________________________

Specimen N: ______________________________________________________________________________

Specimen O: ______________________________________________________________________________

Lab 1 - Woody Twig Terminology 53


Specimen P: ______________________________________________________________________________

Specimen Q: ______________________________________________________________________________

Specimen R: ______________________________________________________________________________

Specimen S: ______________________________________________________________________________

Specimen T: ______________________________________________________________________________

Specimen U: ______________________________________________________________________________

Specimen V: ______________________________________________________________________________

Specimen W: ______________________________________________________________________________

Specimen X: ______________________________________________________________________________

2. In the space below, construct your own dichotomous (winter) key to four woody plant species using only winter
condition twig characteristics. Make sure to use some of the terminology that you are learning in Lab today!

54 Integrative Biology 335: Plant Systematics


3. In the box below, choose one of the twigs you identified, draw it and then label its parts, making sure to include
any diagnostic features (especially those mentioned in the “Dichotomous (Winter) Key”).

Note: The subject of woody twigs in winter condition is covered minimally in lecture. Thus, take some time on your
own to make sure that you understand all of the woody twig terminology. In future labs, as well as on the lab exams,
you may be asked to use a dichotomous key to identify a woody twig.

Lab 1 - Woody Twig Terminology 55


DICHOTOMOUS KEY GUIDELINES

1. Always make sure you’ve selected the appropriate key for the materials to be identified. For example, if the
material to be identified is cultivated, select a manual treating such plants since most floras do not include
cultivated plants unless naturalized.

2. Read the introductory comments on format details, abbreviations, etc. before using the key.

3. Keys are made for average plants. When offered a selection of individual plants avoid the unusual or freakish
specimens. Since living things are always somewhat variable, do not base your conclusions on a single
observation, but arrive at an average by studying several specimens. It is surprising how often students will find
the one unusual or aberrant sample in a large assortment of normal things!

4. Take your time using a key.

5. Always read both choices even if the first seems a logical one to take. The second may be even better. Get
both sides of the picture. The second choice, by contrast, may actually help to explain the first one. Note that in
some keys the author may have used more than two choices. This is disconcerting if the third choice happens to
fall on a different page.

6. Read all the supplemental characters, not just the first ones. However, do not fix on a second or third minor
character and ignore the most important first one.

7. Be sure you understand the meanings of the terms involved. Do not guess.

8. When measurements are given, use a calibrated scale. Do not guess.

9. When minute objects are concerned, use a lens of sufficient magnifying power to show clearly the feature you
need to see.

10. Check yourself at every possible step with the descriptions. For example, when you arrive at a family in the key
(when keying a plant to species) take time to read over the family description before proceeding. This may save
you a lot of trouble later on. Use pictures or drawings when available but remember that a good description is far
superior to a picture in illustrating the range of variation found within the species.

11. If the choice of division is not clear, or if you have no way of making a choice because you do not have sufficient
information, try both divisions, arrive at two possible answers by doing so, and then read descriptions of each in
order to make a choice.

12. Once the material has been keyed out with the “Dichotomous (Winter) Key for Woody Plants in Champaign-
Urbana”, verify your results by reading the description (if available in the literature you are using, or in another
reference which describes the plant), or by comparing the specimen with an illustration or an authentically named
herbarium specimen.

56 Integrative Biology 335: Plant Systematics


DICHOTOMOUS (WINTER) KEY FOR WOODY PLANTS IN CHAMPAIGN-URBANA

1. Leaf scars opposite


2. Bundle scars several and separate, or in several separate groups
3. Leaf scars about as broad as long Aesculus spp.
3. Leaf scars somewhat U- or V-shaped
4. Bud scales overlapping each other, pubescent only on margin Acer rubrum
4. Bud scales not overlapping each other, pubescent throughout Cornus stolonifera
2. Bundle scars single, or closely crowded forming a single circle, ellipse or
crescent
5. Terminal buds absent; twig terminated by a pair of axillary buds Syringa vulgaris
5. Terminal buds present Fraxinus pennsylvanica
1. Leaf scars alternate
6. Plants armed with thorns or spines
7. Armature of thorns in axils of leaf scars Crataegus crus-galli
7. Armature of paired spines at nodes Rosa multiflora
6. Plants unarmed (Note: Mature twigs of Maclura pomifera may be thorny!)
8. Buds crowded at tips of twigs; pith strongly 3-5 sided in cross section
9. Twigs corky ridged Quercus macrocarpa
9. Twigs not corky ridged Quercus rubra
8. Buds distributed rather evenly along twigs, not crowded at tips; pith
circular or not strongly 3-5 sided in cross section
10. Bundle scars single, or closely crowded forming a single circle, ellipse
or crescent
11. Stipule scars present Maclura pomifera
11. Stipule scars absent Diospyros virginiana
10. Bundle scars several and separate, or in several separate groups
12. Stipule scars present
13. Stipule scars encircling twig completely or nearly so
14. Stipule scars completely encircling twig
15. Buds encircled by leaf scars Planatus occidentalis
15. Buds not encircled by leaf scars
16. Leaf scars nearly circular-shaped; buds not Liriodendron tulipifera
silky
16. Leaf scars crescent-shaped; buds densely silky Magnolia acuminata
14. Stipule scars nearly encircling twig and slightly offset Fagus sylvatica
13. Stipule scars extending less than halfway around twig
17. Bud scales one per bud Salix spp.
17. Bud scales more than one per bud
18. Buds with 2 or 3 visible bud scales Tilia americana
18. Buds with more than 3 visible bud scales
19. Leaf scars in more than 2 ranks; bark with Prunus serotina
flavor and odor of almonds when crushed
19. Leaf scars in 2 ranks; bark not as above Betula pendula
12. Stipule scars absent
20. Pith chambered Juglans nigra
20. Pith of uniform texture
21. Axillary buds nearly or completely surrounded by leaf
scars
22. Twigs densely velvety-hairy Rhus typhina
22. Twigs smooth Cladrastis lutea
21. Axillary buds not surrounded by leaf scars
23. Bundle scars many; leaf scars large and circular- Carya ovata
shaped
23. Bundle scars 3; leaf scars small and broadly Liquidambar styraciflua
crescent-shaped

Lab 1 - Woody Twig Terminology 57


BOOK RESOURCES

Several or all of the following books are on display in the lab and have keys to woody plants in winter condition.

1. Barnes, B.V. and W.H. Wagner, Jr. 1981. Michigan Trees. The University of Michigan Press, Ann Arbor, Michigan.

2. Campbell, C.S. and F. Hyland. 1975. Winter Keys to Woody Plants of Maine. University of Main Press, Orono,
Maine.

3. Core, E.L. and N.P. Ammons. 1958. Woody Plants in Winter. The Boxwood Press, Pacific Grove, California.

4. Harlow, W.M. 1941. Fruit Key and Twig Key to Trees and Shrubs. Dover Publications Inc., New York.

5. Hosie, R.C. 1969. Native Trees of Canada. Canadian Forestry Service, Department of the Environment, Ottawa,
Ontario, Canada.

6. Preston, R.J., Jr. and V.G. Wright. Identification of SE Trees in Winter.

7. Trelease, W. 1931. Winter Botany. Urbana, Illinois.

8. Viereck, L.A. and E.L. Little, Jr. 1972. Alaska Trees and Shrubs. Forest Service, U.S.D.A., Agriculture Handbook
No. 410.

ACKNOWLEDGEMENTS

Much of the material presented in this lab exercise was taken from handouts Dr. Downie used while he was an
undergraduate student at the University of Guelph, Ontario, Canada. This introductory plant taxonomy course was
taught by Dr. Judith Canne-Hilliker. Other than that, the source of the material is unknown.

58 Integrative Biology 335: Plant Systematics


GLOSSARY OF WOODY TWIG TERMINOLOGY

Listed below is the woody twig terminology that you will need to become skillful at using this semester. You should be
able to draw, compare and contrast each of these terms.

WOODY TWIG PARTS

TWIG
The terminal portion of a branch of a woody
plant. In most keys this term is used to
designate, specifically, the growth of the
current year and, to a lesser extent, that of the
last preceding year.

NODE
The position on a stem where a leaf or bud is
(or was) attached.

INTERNODE
The portion of a stem between two nodes.

AXIL
The upper angle between a leaf (or any other
lateral structure) and the stem to which it is
attached.

BUDS
The early undeveloped stage of a leafy shoot
or flower. These growing points are dormant in
winter and are usually covered, for protection,
by bud scales.

BUD SCALES
Modified leaves (or, rarely, stipules) that serve
to protect the bud.

BUD SCALE SCARS


A ring of scars around the twig that marks
the place where growth began that year. The
distance from the terminal bud to the first ring
of bud scale scars represents the amount of
growth in length that occurred during the most
recent growing season.

TERMINAL BUDS
Buds formed at the tips of the stem and its
branches.

AXILLARY (LATERAL) BUDS


Buds formed alongside the twigs and arising in
the axils of the leaves. Often more than one
bud appears at a node, in which case the one
directly above the leaf scar is considered the
true axillary bud and the others are designated
as accessory or collateral buds.

Lab 1 - Woody Twig Terminology 59


WOODY TWIG PARTS (cont.) PITH TYPES [TWIG L.S.]

PITH CONTINUOUS PITH


The central portion of a twig. Solid pith (no partitions or chambers)

LEAF SCARS DIAPHRAGMED PITH


The place from which the leaf has fallen. This Solid pith with partitions
region differs in color and texture from the rest
of the twig. Shapes of scars are quite variable. CHAMBERED PITH
Hollow pith with partitions
BUNDLE SCARS
These indicate the broken ends of the vascular
bundles passing from the stem into the leaves.
They are identified as small dots or lines on the
surface of the leaf scar. Also called “Traces”.

STIPULES
A pair of appendages at the base of a leaf stalk
(petiole) and often adnate to it. Stipules are
usually leaf-like but can be modified in some
plants into prickles or tendrils.

STIPULE SCARS
CONTINUOUS PITH DIAPHRAGMED PITH CHAMBERED PITH
These are generally slit-like in shape and
inconspicuous. In a few species (e.g.
Liriodendron tulipifera) they encircle the twig
from one edge of the leaf scar to the other.
PITH SHAPES [TWIG X.S.]

SPECIAL WOODY TWIG PARTS


ROUND 3-ANGLED 5-ANGLED 5-POINTED
LENTICILS
Small, often wart-like prominences scattered
over the surface of twigs. They serve to admit
air to the living tissues beneath.
LEAF SCAR SHAPES
SPUR SHOOTS
Short, stubby branches with greatly crowded
leaf scars and very slow growth (e.g. Betula
spp.). Often
flower buds HORSESHOE-SHAPED U-SHAPED V-SHAPED
are produced
on short
shoots (e.g.
Malus spp.)
CRESCENT-SHAPED BROADLY CRESCENT SHIELD-SHAPED

HALF-ROUND ROUNDED OR CIRCULAR OVAL

TRIANGULAR THREE-LOBED HEART-SHAPED

60 Integrative Biology 335: Plant Systematics


BUNDLE SCAR ARRANGEMENT LEAF AND SCALE SCARS

STIPULES AND STIPULE SCARS

Lab 1 - Woody Twig Terminology 61


62 Integrative Biology 335: Plant Systematics
LABORATORY 2
VEGETATIVE TERMINOLOGY

Phytography is that part of systematics which deals with descriptions of plants and their various organs, such as
roots, stems, and leaves. The variation in size, shape, texture, and structure of these organs provides data that
taxonomists use in the naming, identification, and classification of plants. In this class we provide you with a list of
terms that describes some of this variation. These terms were selected for you to know because of their widespread
use in taxonomy. In this Lab you will become acquainted with some of the terminology used to describe the
vegetative body of vascular plants. By working through this exercise and studying the plants carefully, you should
be able to obtain a good working knowledge of these basic terms. Enjoy the plants and our wonderful greenhouse
facilities!

TAXONOMY TERMS

Taxonomic characters are fundamental to the science of taxonomy. They provide the basic information for
classification, they are the features used in identification, they are used to determine relationships, and they give
the basis for the naming of taxa. You should know the definition of a TAXONOMIC CHARACTER as well as be able to
distinguish between CHARACTER and CHARACTER STATES.

TAXONOMIC CHARACTER
Any attribute of an organism that can be consistently measured, counted, described, or otherwise assessed (e.g.
leaf shape, plant height, stamen number, pigment type, DNA nucleotide)

CHARACTER
A group of states considered to be modifications or CHARACTER CHARACTER STATE
alternate forms of basically the same thing.
Flower Color Red, white or blue
CHARACTER STATE
Leaf Shape Linear, lanceolate or reniform
One of the various conditions (attributes, or values) of
a character observed across a group of taxa. It is the DNA nucleotide A, C, G or T
fundamental unit of a CHARACTER.

LAB EXERCISE

During today’s Lab in the Plant Sciences Laboratory, you will practice using vegetative terminology to describe
flowering plants. The TA will begin by giving you a brief orientation to the Conservatory and the teaching collection
rooms 1401 - 1415. Afterward, you will complete the following Lab exercise by visiting each room, examining the
plants listed and answering the questions with the aid of the Glossary of Vegetative Terminology provided at the end
of the Lab. To avoid conjestion, you may begin in any room. Just a few reminders: Handle the plants carefully! You
may touch them, but please do not destroy them. Also, walk only on the walkways in the Conservatory and make sure
to close all doors behind you after entering and exiting a each room (most of the collection rooms are temperature
controlled.)

Lab 2 - Vegetative Terminology 63


PLANT BIOLOGY CONSERVATORY AND COLLECTION ROOMS

The Conservatory and Plant Collection rooms were created in 1988. By September 1991 the Conservatory landscape
installation was finished, including a small pond and waterfall. In total, these facilities occupy 4,000 square feet
of the Plant Sciences Laboratory building and house over 1,100 species from 161 familes. While walking through
the Conservatory, you will see over 200 plant species from about 70 plant families. Most of these species are
representative of tropical rainforests, and some are economically important. From trees and shrubs to ferns and
vines, you will also see many plant forms. Each Plant Collection room showcases plants from a unique habitat type.

A NOTE ON NOMENCLATURE

Cultivated varieties of plants, or cultivars, are preceded by the letters “cv.” or their names are included in single
quotation marks. The cultivar name is not italicized. For example, both of the following names are written correctly
and refer to exactly the same cultivar:

Begonia rex ‘Fireworks’


Begonia rex cv. Fireworks

ROOM 1201 - THE CONSERVATORY

Cycas circinalis L. (CYCADACEAE) (Queen Sago)

Cycads, or tropical evergreens, were the dominant flora in the early


Mesozoic (200 million years ago). C. circinalis, one of the six species
of cycads found in the Conservatory, is DIOECIOUS. What does that
mean? Make sure to check out its palm-like leaves and the LEAF SCARS
that cover its trunk. Be aware that Cycas is a gymnosperm, not an
angiosperm.

Amorphophallus titanum (Becc.) Becc. (ARACEAE) (Titan Arum)

The titan arum (or corpse flower) develops from a CORM and then
spends years (6 - 7) in the vegetative part of its lifecycle. (Fun Fact:
CORMS from this species can be over 200lb!) What is a CORM?

After the vegetative stage is complete, the species will bloom and emit
a “rotting-fish-with-burnt-sugar” odor to attract its natural pollinators,
carrion beetles and flesh flies. The titan arum is often said to have the
“world’s largest flower”. Is this a botanically accurate statement given
that this species is in the same family as Anthurium? Why or why not?

64 Integrative Biology 335: Plant Systematics


Musa spp. (MUSACEAE) (Banana)

This economically important plant grows from an underground stem


(RHIZOME) and its berry fruit is sterile, so no seeds are produced. Why
do we consider this plant a HERB and not a TREE in the true “botanical”
sense of the word? What is its LIFE SPAN? LEAF VENATION?

Oxalis spp. (OXALIDACEAE) (Oxalis)

This genus belongs to the wood-sorrel family and is characterized by


having palmately compound leaves typically composed of three to ten
leaflets. What is the special LEAF COMPLEXITY term applied to this leaf
since it has three leaflets? What is the BLADE SHAPE of the leaflets?

Theobroma cacao L. (STERCULIACEAE) (Chocolate Tree)

For fellow chocolate lovers like myself, it will come as no surprise that
the genus name for this plant, Theobroma, is Greek for “food of the
gods”. T. cacao are small EVERGREEN trees that produce fly-pollinated
flowers directly on their branches. The 20-60 seeds found in each
mature fruit is what is used to produce chocolate. Make sure to look at
the flowers and pods on this individual!

Barringtonia racemosa Roxb. (LECYTHIDACEAE) (Powderpuff Tree)

B. racemosa is aptly named


since its flowers hang in long
racemes. Make sure to observe
the flowers’ prominent stamens
in preparation for our floral
morphology lab next week.
Common along the Indian
Ocean’s tropical coasts, its
fruit have evolved to be very
buoyant so that they can be
dispersed by water. What
would the PLANT HABIT and
LEAF APEX be classified as for
this species? Draw the LEAF
APEX in the box provided.

Lab 2 - Vegetative Terminology 65


Anthurium spp. hybrid (ARACEAE) (Flamingo flower)

Characteristic of the Araceae


family, this plant’s inflorescence
is composed of tightly packed
flowers arranged on a spadix.
Surrounding the spadix is a
colorful bract called a spathe.
Observe the leaves for
Anthurium. Sketch and label
the LEAF SHAPE and MARGIN in
the box provided.

ROOM 1415 - TROPICAL PLANTS

Streptocarpella saxorum Engler (GESNERIACEAE) (Cape Primrose)

In order to propagate S. saxorum, one must divide the root ball or grow
leaf cuttings since the flowers are typically sterile. As you can see, the
flower petals of this species are fused (connate) and can be described
as sympetalous, but more about that next week. What terms describe
the PLANT HABIT and the LEAF SURFACE?

Callisia fragrans (Lindl.) Woodson (COMMELINACEAE) (Basket Plant)

Not only is C. fragrans a decorative plant, but it is also a widely used


in Russia for medicinal purposes. What is its LEAF VENATION and STEM
TYPE?

Salvia leucantha Cav. (LAMIACEAE) (Mexican Bush Salvia)

Look for the distinctive


STIPULES on S. leucantha.
Name the LEAF ARRANGEMENT
of this species and sketch it in
the box provided.

66 Integrative Biology 335: Plant Systematics


Mimosa pudica L. (FABACEAE) (Sensitive Plant, Shameplant)

The species name, “pudica” is very appropriate for this plant in that it
means bashful. Touching the leaves of M. pudica stimulates it to release
chemicals, thereby causing the redistribution of water in the cells. The
unequal distribution of water is what allows the leaves to fold up. This
mechanism is speculated to be a defense against herbivory.

Cissus rotundifolia (Forssk) Vahl (VITACEAE) (Venezuelan Treebine)

Describe the LEAF BLADE SHAPE of C. rotundifolia. Its specific epithet


should give you a clue! Also, what is the name of the twining structure
that this plant uses to climb or support itself?

Schefflera arboricola (Hayata) Kanehira (ARALIACEAE)


(Dwarf Umbrella Tree)

This EVERGREEN
shrub is native to
Taiwan and is often
used as a houseplant.
What is the LEAF
COMPLEXITY of this
plant? Draw one leaf
in the box provided
and label its parts.

ROOM 1413 - EPIPHYTIC PLANTS, ORCHIDS AND BROMELIADS

Miscellaneous spp. (ORCHIDACEAE)

In many Orchidaceae species, the leaves are attached to a swollen


fleshy stem called a PSEUDOBULB. What do you think is the purpose
of a PSEUDOBULB? While in this room make sure to check out all the
blossoms! We’ll be studying Orchidaceae flower structure later in the
semester.

Lab 2 - Vegetative Terminology 67


Phalaenopsis hybrid (ORCHIDACEAE)

Epiphytic Orchidaceae, such as this Phalaenopsis hybrid, are often


characterized by aerial, cord-like roots. These roots are adventitious and
are covered by a multi-layered epidermis (called VELAMEN) that absorbs
moisture from the atmosphere.

Vanilla planifolia Jacks. (ORCHIDACEAE) (Flat-leaved Vanilla)

Consider V. planifolia, the source of the flavoring vanilla (from the cured
fruits or pods). This species was used by the Aztecs to flavor cocoa.
Once more note the large aerial, adventitious roots. What is the LEAF
ARRANGEMENT and PLANT HABIT?

Dendrobium nobile ‘Montrose’ (ORCHIDACEAE)

Notice the conspicuous parallel leaf venation on D. nobile, which is


characteristic of Orchidaceae and monocots in general. Classify the
LEAF ATTACHMENT.

Vriesea carinata Wawra (BROMELIACEAE) (Painted Feather)

Bromeliads, such as V. carinata, are acaulous herbs (the prefix “a” =


without; “caulous” mean stem). Their leaves are usually leathery or
fleshy, covered in a thick cuticle and clustered in a tight rosette. Thus,
the LEAF ARRANGEMENT could be called what?

Ananas comosus (L.) Merr. (BROMELIACEAE) (Pineapple)

A. comosus produces a multiple fruit of berries, commonly known as


a pineapple. Generally hummingbirds are the main pollinators of this
species but are purposefully kept away from pineapple plantings to
avoid the development of seeds in the fruit. Most pineapple plants are
propagated by vegetative means. Describe the LEAF SHAPE and LEAF
APEX.

68 Integrative Biology 335: Plant Systematics


ROOM 1411 - SUBTROPICAL PLANTS AND CACTACEAE

Rhipsalis cereuscula Haworth (CACTACEAE) (Coral Cactus)

The cacti in this room are typical of tropical environments. In particular,


R. cereuscula is an epiphytic cactus that has jointed stems and
spherical translucent and mucilaginous fruits. Can you describe its
HABIT?

Carissa macrocarpa (Eckl.) A. DC. (APOCYNACEAE) (Natal Plum)

This plant produces plum shaped fruits that are edible when ripe,
however don’t try to eat any other part of C. macrocarpa – it’s
poisonous! What terms describe its LEAF APEX and LEAF SURFACE?

Piper auritum Kunth (PIPERACEAE) (Root Beer Plant)

Crush a small bit of the leaf the TA has pulled off and smell it. P. auritum
is often used as a flavoring due to its spicy aromatic scent and flavor.
How would you classify its LEAF ARRANGEMENT and LEAF BASE?

Helwingia chinensis Batalin (HELWINGIACEAE)

The genus containing these deciduous shrubs, Helwingia, was named


for the German botanist Georg A. Helwing. What is H. chinensis’s LEAF
SHAPE and LEAF MARGIN?

Fatsia japonica (Thunb) Decne & Planch (ARALIACEAE) (Paperplant)

Classified as an evergreen
shrub, F. japonica is often used
for landscaping purposes.
What is F. japonica’s LEAF
VENATION and LEAF MARGIN?
Also, how would you classify
the LEAF ATTACHMENT? Draw
and label a leaf in the box
provided.

Lab 2 - Vegetative Terminology 69


ROOM 1409 - DESERT PLANT COLLECTIONS

Opuntia brasiliensis (Willd.) Haw. (CACTACEAE) (Tree Opuntia)

Here is a plant trying to pretend it has broad, flat leaves when in


actuality it has branching stems. What is the plant’s HABIT? Though this
species of Opuntia is native to Brazil, there are three other members of
its genus that are native to Illinois (O. humifusa, O. macrorhiza and O.
fragilis (Endangered in IL)).

Cephalocereus senilis (Haw.) Pfeiff. (CACTACEAE) (Old-Man Cactus)

C. senilis is actually rare and endangered in the wild, however its


appearance has made it commercially popular and thus widespread in
commercial nurseries. Why do you think this plant has evolved to be
covered in grayish-white hairs? In any case, doesn’t it look like Cousin It
from the Addams Family?

Fouquieria fasciculata Nash (FOUQUIERIACEAE) (Árbol de Barril)

The leaves of F. fasciculata are DECIDUOUS but a part of the LEAF


PETIOLE can harden to form the SPINES you see covering most of the
plant. What is the LEAF ARRANGEMENT?

Welwitschia mirabilis Hook. (WELWITSCHIACEAE) (Tree Tumbo)

W. mirabilis is a DIOECIOUS PERENNIAL and the only species in the


Welwitschiaceae family. Make sure to read the plaque posted – it
contains a lot of interesting information about this bizarre species! What
ROOT TYPE does it have? Be aware that Welwitschia is a gymnosperm,
not an angiosperm.

Lithops spp. (AIZOACEAE) (Stone Plant)

These unique plants are made up of one pair of SUCCULENT leaves,


highly modified to retain moisture in times of extreme heat and drought.
Their genus name, Lithops, comes from the Greek “lithos” (a stone) and
“ops” (like). Neat aren’t they?

70 Integrative Biology 335: Plant Systematics


Alluaudia ascendens Drake (DIDIEREACEAE) (Fantsiolotse)

This DIOECIOUS, DECIDUOUS tree has vertical pairs of leaves arranged


under each SPINE. What is the difference between a SPINE, THORN and
PRICKLE?

Bulbine alodies (LILIACEAE)

The species name for B. alodies translates to “aloe-like”. This plant,


unlike Opuntia brasiliensis, is a LEAF SUCCULENT. Be sure you can
distinguish between a LEAF SUCCULENT (such as those belonging
to families Agavaceae, Crassulaceae and Liliaceae) and a STEM
SUCCULENT (i.e. Cactaceae).

ROOM 1405 - CARNIVOROUS AND AQUATIC PLANTS

Marsilea mutica Mett. (MARSILEACEAE) (Floating Water Clover)

This plant is not an angiosperm (it’s a fern actually) but check out its
COMPOUND LEAF with four PINNAE. Is this the quintessential four-leaf
clover? Make a wish!

Spiranthes odorata (Nutt) Lindl (ORCHIDACEAE) (Lady’s Tresses)

Marsh Lady’s tresses is considered an endangered orchid species in


three states (Kentucky, Maryland and Tennessee). Make sure to take a
look at its beautiful, white flowers! What is its HABIT, LEAF ARRANGEMENT
and LEAF VENATION?

Cyperus papyrus L. (CYPERACEAE) (Papyrus Plant)

C. papyrus was the source of papyrus more that 4,000 years ago! The
stem pith was sliced into thin strips, then laid side by side. Another
layer of strips was added over the top at right angles and the two layers
were hammered together until a sheet of “paper” was formed. Note the
triangular stems, 3-ranked leaves and closed sheaths (the Cyperaceae
family is characterized by these features).

Lab 2 - Vegetative Terminology 71


Colocasia esculenta (L.) Schott (ARACEAE) (Coco Yam, Taro)

This large herb arises from an underground TUBER. Describe the LEAF
BASE (Hint: Where does the stem attach?) Also, what type of LEAF
VENATION does this plant have and is this typical of the Araceae family
(a monocot family)?

CARNIVOROUS PLANTS

Nepenthes spp. (NEPENTHACEAE) (Pitcher Plants)

These plants possess hollow, tubular leaves which are urn- or trumpet-
shaped and often resemble small pitchers. Note the TENDRIL at the tip
of each leaf. Insects are attracted by nectar secretions near the rim and
when they fall in, the sharp, downward pointing hairs within the pitcher
prevent the insects’ escape. Digestive enzymes secreted by internal
glands then digest the insect.

Drosera spp. (DROSERACEAE) (Sundews)

Drosera’s LEAF SURFACE can be described as GLANDULAR due to the


stalked glands found on on their often club-like or linear-shaped leaves.
These glands secrete a sticky liquid that catches the prey. Enzymes are
also secreted that dissolve the victim, and the resultant nutrients are
absorbed by numerous tiny hairs on the leaf.

Dionaea spp. (DROSERACEAE) (Venus Flytrap)

In these plants the leaves are divided into a LEAF BLADE and a
predominantly winged PETIOLE. The blade is made up of two identical
semicircular halves united along the middle by a thick midrib. Together
they constitute, in effect, a hinged trap. The outer margin of each lobe
is characterized by a comb of long, bristle-like teeth. Yes, the archetypal
man-eating plant!

ROOM 1403 - ECONOMIC PLANTS

Ficus carica L. (MORACEAE) (Edible Fig)

Ficus species are characterized by their “fruit-like” structures called


syconia, or figs, that contain many small male and female flowers. Many
Ficus species require a wasp to enter the syconium to pollinate its
flowers, however F. carica does not. This is fortunate for oftentimes the
wasps will die inside the synconium. Make sure to check out the LEAF
SCARS on F. carica’s branches. What is its LEAF MARGIN?

72 Integrative Biology 335: Plant Systematics


Manihot esculenta Crantz (EUPHORBIACEAE) (Cassava)

The TUBEROUS ROOT from M. esculenta is one of the world’s major


food crops. The original bitter variety of cassava contained cyanide
compounds and thus was poisonous unless they were peeled and
boiled multiple times. However, new sweet varieties have been bred
so that they can be eaten raw when peeled (the cyanide compounds
still remain in the TUBER’s skin regardless of the variety). What is M.
esculenta’s LEAF ARRANGEMENT and LEAF VENATION?

Mentha suaveolens Ehrh. (LAMIACEAE) (Wooly Apply Mint)


Origanum vulgare L. (LAMIACEAE) (Greek Oregano)

Look, touch, feel and smell these two plant species. What do they have
in common? Hint: What is their HABIT? LEAF ARRANGEMENT? Shape of
the stem in cross-section? These features are all characteristic of the
Lamiaceae family. The odor comes from small glands on the leaves and
stems that contain ethereal oils.

Cinnamomum zeylanicum Garcin (LAURACEAE) (Cinnamon)

C. zeylanicum is one of four Cinnamomum species that are utilized for


the commercial cultivation of cinnamon. To produce cinnamon, these
trees are cut at ground level and the inner bark is removed in rolls and
cut into pieces (cinnamon sticks). What terms describe C. zeylanicum’s
LEAF SHAPE, MARGIN and ARRANGEMENT?

Swietenia macrophylla King (MELIACEAE) (Big Leaf Mahogany)

S. macrophylla has been


placed on the endangered
species list in its native
region and thus is no longer
harvested there for its beautiful
wood. It has been introduced,
however, in other tropical areas
for commercial production
(in some cases it’s actually
become invasive!). What is
the LEAF COMPLEXITY of S.
macrophylla? Draw and label a
leaf in the box provided.

Lab 2 - Vegetative Terminology 73


Coffea arabica L. (RUBIACEAE) (Arabian Coffee)

If you’re a crazy coffee drinker like your TA, you’ll appreciate this
amazing plant. (Fun Fact: Americans drink 400 million cups of coffee per
day.) Check out the red fruits (drupes) that each contain two seeds, or
“beans”. In order to produce the best coffee, the fruits need to be hand
harvested when they deep red color - not before or later. Determine the
LEAF ARRANGEMENT and LEAF SURFACE of the plant.

Saccharum officinarum L. (POACEAE) (Sugar Cane)

This is sugar cane, the source of half the world’s sugar. Look closely at
its stem. What is the LEAF ATTACHMENT and LEAF VENATION?

74 Integrative Biology 335: Plant Systematics


VEGETATIVE WORKSHEET

Choose one plant in the Conservatory or the teaching collection rooms and complete the following worksheet using
the terminology covered in today’s Lab. (By describing all of your selected plant’s character states, you are essentially
creating a species description.) Then in the box below draw the species or at least one of its key features.

FAMILY ______________________________________ LEAF (OR LEAFLET) BLADE SHAPE __________________

SPECIES _____________________________________ LEAF VENATION ________________________________

CULTIVAR (IF APPROPRIATE) _______________________ LEAF MARGIN _________________________________

PLANT HABIT _________________________________ LEAF APEX ___________________________________

LIFE SPAN ____________________________________ LEAF BASE ___________________________________

STEM TYPE ___________________________________ LEAF SURFACE ________________________________

LEAF COMPLEXITY _____________________________ STIPULES (PRESENT OR ABSENT) ___________________

LEAF ARRANGEMENT ____________________________ OTHER SPECIAL FEATURES ________________________

LEAF ATTACHMENT ______________________________ _____________________________________________

NOTES _______________________________________________________________________________________

_____________________________________________________________________________________________

_____________________________________________________________________________________________

Lab 2 - Vegetative Terminology 75


GLOSSARY OF VEGETATIVE TERMINOLOGY

Listed below is the vegetative terminology that you will need to become skillful at using this semester. You should be
able to draw, compare and contrast each of these terms. You should also know the relationship between each of the
terms listed (i.e. CHARACTER STATES) and the major category (i.e. CHARACTER) to which they belong.

A Few Words of Caution: Terms were coined to represent clearly distinguishable differences. However, plants
vary continuously, and the structure that you see on a particular plant may not precisely match the illustrations or
definitions provided in this Lab Manual or in the Course Textbook. Often you will encounter a feature that seems to be
intermediate between two terms (e.g. a leaf that is between lanceolate and broadly ovate in shape). As you become
more familiar with a specific feature, you will gradually develop an understanding of the range of variability that it
encompasses.

PLANT HABIT

HERB
No above ground persistent woody tissue but may
have underground perennating structures

SUB-SHRUB
Lower stems woody but upper stems herbaceous

SHRUB
A woody low-stature perennial plant with one to SUCCULENT (E.G. STEM)
many slender trunks arising from near its base
HERB
TREE
A large woody perennial plant with one to several
relatively massive trunks and an elevated crown

SUCCULENT
Possessing thick, usually soft, watery leaves and/
or stems (there are stem & leaf succulents)

VINE
A woody or herbaceous plant with a long, slender,
more or less flexible stem which cannot support
itself
LIANA
LIANA
A woody, climbing vine (characteristic of the
tropics)

VINE SHRUB TREE

76 Integrative Biology 335: Plant Systematics


LIFE SPAN (DURATION) STEM TYPE

ANNUAL AERIAL
Completes life cycle in one growing season An erect or prostrate stem (the most common)

BIENNIAL STOLON
Completes life cycle in two growing seasons A horizontal stem near the ground surface that
produces new plants at its nodes or tip (runner)
PERENNIAL
Lives for more than two growing seasons RHIZOME
An underground horizontal persistent stem;
its leaves are often reduced to scales and it
usually bears adventitious roots and buds
LEAF PERSISTENCE
BULB
A thickened, underground, short, vertical stem
DECIDUOUS with large storage leaves; usually below ground
Loses leaves during unfavorable conditions
(such as at the end of each growing season) CORM
A solid, erect, underground stem with leaves
EVERGREEN absent or dry and scale-like
Bearing green foliage all year round
TUBER
A solid, enlarged, underground, horizontal stem
that serves as a storage area for food reserves
ROOT TYPE

TAPROOT
Central main root that descends vertically;
larger than any branching root

FIBROUS
Thin, thread-like roots arising from a taproot or
from stem tissue
AERIAL STOLONS

ADVENTITIOUS
Roots that originate from any part of the plant
other than the root system

RHIZOME BULB

TAPROOT FIBROUS ADVENTITIOUS

CORM TUBER

Lab 2 - Vegetative Terminology 77


LEAF COMPLEXITY

To determine the complexity of a leaf, always


APEX
look for the AXILLARY BUD located above the LEAFLET
PETIOLE. Anything above the AXILLARY BUD is (PINNA)

considered one leaf. LEAF BLADE

A SIMPLE LEAF will have a single blade attached


to the petiole, whereas a COMPOUND LEAF will
have more than one blade attached to the RACHIS
petiole.
MARGIN
PETIOLULE
VENATION

BASE PETIOLE
SIMPLE LEAF
AXILLARY BUD
A leaf with a single blade per petiole (it is not
divided into leaflets; there is always a flange of
SIMPLE LEAF COMPOUND LEAF
blade tissue connecting adjacent lobes)

TRIFOLIATE
Three separate leaves arising from the
same node

COMPOUND LEAF
A leaf with more than one blade per petiole (it
TRIFOLIATE
is made up of two or more leaflets and these
leaflets are wholly separate) TRIFOLIOLATE

TRIFOLIOLATE
A compound leaf with three leaflets

PINNATELY COMPOUND LEAF


A leaf in which there are more than three
leaflets (PINNA, pl. PINNNAE) arranged in
two rows along a 1° rachis.
PALMATELY COMPOUND
BIPINNATELY COMPOUND LEAF
A leaf that is divided twice (has a 1° and 2° PINNATELY COMPOUND
rachis)

TRIPINNATELY COMPOUND LEAF


A leaf that is divided three times (have a
1°, 2° and 3° rachis)

PALMATELY COMPOUND LEAF


A compound leaf where the leaflets arise
from a common point of attachment (no
rachis)

BIPINNATELY COMPOUND TRIPINNATELY COMPOUND

78 Integrative Biology 335: Plant Systematics


LEAF PARTS LEAF ARRANGEMENT

LEAF BLADE BASAL


Expanded portion of a leaf Leaves attached to a rhizome or underground
stem
LEAFLET
One of the segments of a compound leaf CAULINE
Leaves attached to an above-ground stem
PINNA (pl. PINNNAE)
The leaflets of a pinnately compound leaf ALTERNATE
One leaf per node
PINNULE
The leaflets of a bipinnately and tripinnately OPPOSITE
compound leaf Two leaves per node

PETIOLE WHORLED
Stalk of a leaf Three or more leaves per node

PETIOLULE
The stalk of a leaflet (i.e. PINNA or PINNULE)

RACHIS
The axis of a pinnately compound leaf.

BASAL CAULINE
LEAF ATTACHMENT

PETIOLATE
Leaf has a petiole

SESSILE
Leaf lacks a petiole

SUBSESSILE
Leaf posseses a very short petiole

SHEATHING
Leaf base enwraps stem

ALTERNATE OPPOSITE WHORLED

PETIOLATE SHEATHING

SESSILE

Lab 2 - Vegetative Terminology 79


LEAF BLADE SHAPE

LINEAR OBOVATE
Long and narrow with the side parallel (>4:1) Ovate, but with narrower end towards point of
attachment (The prefix “ob” means opposite, so
OBLONG “obovate” is the opposite of “ovate”)
Nearly rectangular with the sides parallel (2-
4:1) ELLIPTIC
Widest near middle and tapering at both ends
LANCEOLATE
Spear-shaped; widening above base and then ORBICULAR
long tapering to apex (3-4:1) Circle shaped

OVATE RENIFORM
Egg-shaped; broadest near base (<3:1) Kidney shaped

LINEAR OBLONG LANCEOLATE OVATE OBOVATE ELLIPTIC ORBICULAR RENIFORM

LEAF VENATION LEAF APICES

PARALLEL ACUMINATE
Veins lie roughly parallel to the leaf margins. Sharp, ending in a long-tapering point with
concave sides
PINNATE
Central mid-vein with many 2° veins emerging ACUTE
on both sides. Sharp, ending in a point with straight sides to
the apex (< 90°)
PALMATE
All 1° veins arise at the same point at the base OBTUSE
of the leaf. Blunt, rounded (> 90°)

MUCRONATE
A small, abrupt point

PARALLEL PINNATE PALMATE ACUMINATE ACUTE OBTUSE MUCRONATE

80 Integrative Biology 335: Plant Systematics


LEAF BASES LEAF MARGIN

ACUMINATE ENTIRE
Sharp, long-tapering point A margin without any toothing or division

ACUTE SERRATE
Sharp (< 90°) Sharp teeth pointing toward the apex

OBTUSE CRENATE
Blunt (> 90°) Scalloped or round-toothed

CORDATE DENTATE
Heart-shaped (equal rounded lobes at the Sharp teeth projecting at right angles from the
base) margin

OBLIQUE PINNATELY LOBED


Unequal sized lobes at base Lobes towards the midrib but not reaching it

PELTATE PALMATELY LOBED


Umbrella like; the petiole is attached to the Lobes all arising from one point at the leaf base
blade inside of the margin; leaf often orbicular
in shape

ACUMINATE ACUTE OBTUSE ENTIRE SERRATE CRENATE DENTATE

CORDATE OBLIQUE PELTATE PINNATELY LOBED PALMATELY LOBED

LEAF SURFACE

GLABROUS
Lacking hairs (TRICHOMES); a smooth surface

GLANDULAR
Hairs that bear glands that break down into sticky GLABROUS GLANDULAR

beads of fluid; may be stalked (STIPITATE) or sessile

PUBESCENT
Covered with hairs (TRICHOMES)
PUBESCENT STELLATE
STELLATE
Hairs that branch at or near their base (star-shaped from above)

Lab 2 - Vegetative Terminology 81


SPECIAL FEATURES

PRICKLE
A sharp-pointed outgrowth from the epidermis

SPINE
A sharp-pointed modified leaf or leaf part

STIPULAR SPINE
Borne in pairs and lateral to leaf (or leaf scar)

TENDRIL
An elongated, twining segment of a leaf, stem
or inflorescence by which a plant clings to its
support

THORN
A woody, sharp pointed, modified stem (has
stem-like vasculature)

PRICKLE SPINE

STIPULAR SPINE

TENDRIL THORN

82 Integrative Biology 335: Plant Systematics


LABORATORY 3
FLORAL AND INFLORESCENCE TERMINOLOGY

“If I could paint the flower exactly as I see it no one would see what I see because I would paint it small like the flower
is small. So I said to myself... I’ll paint it big.” - Georgia O’Keefe, 1939

The flowering plants are the most numerous, most diverse and most important plants over much of the earth’s
land surface. One characteristic feature of the angiosperms is the grouping of reproductive structures with sterile
auxiliary ones into a single unit known as the flower. An understanding of their reproductive structures is basic to any
introductory taxonomy course. At the end of this Lab exercise you will find an illustrated glossary containing many of
the floral terms you must learn.

FLORAL DRAWINGS

As part of all future Lab exercises, you


will be required to make floral drawings. Species Name Floral Formula Your Name
For some of you, this will be easy and (Family Name) Lab Number
perhaps even fun. For others, like your Date
professor, who have no natural drawing
abilities, this will be laborious at best.
However, experience has shown that
this is the best way to get you to look L. S. of Flower
STIGMA
at the material closely, to understand GYNOECIUM
STYLE
the important points, and to record your (CARPELS)
OVARY
observations for later reference (such
as studying for an exam.) Your drawings
will be turned in and graded.

The important point to remember


ANTHER
is that the artistic quality of the ANDROECIUM COROLLA (PETALS)
drawings is not of any consequence. (STAMENS)
What matters is that the structures FILAMENT
OVULE
are recorded accurately and in their PERIANTH
proper relationship. “Stick drawings” or
CALYX (SEPALS)
diagrams that do this will be graded just
as highly as drawings that look like they
were drawn by Leonardo da Vinci. In PEDICEL RECEPTACLE

the same light, detailed drawings that


are artistically beautiful but botanically
incorrect will be given low grades.

The most common drawings will be of X. S. of Ovary


flowers, especially median longitudinal
sections and cross sections of ovaries. OVARY WALL
An example drawing is shown to the
right. Please mimic the format presented OVULE
SEPTUM
in this example EXACTLY. If you have
questions about your drawings, please
LOCULE
see your TA or your professor!

Lab 3 - Floral and Inflorescence Terminology 83


For full points on your floral drawings, be sure to:

1. Make high quality flower dissections and understand the structure


before drawing. Check with the TA if you have questions!

2. Use a quality pencil and have an eraser.

3. Include the scientific name (underlined!) and family.

4. Write out the complete floral formula. This should be of the plant
observed, not necessarily the general floral formula for the family.

5. Include your own name, date and lab section.

6. Make your drawings large enough so that the important details can
be seen.

7. Label all important parts in the drawings.

8. Use a ruler to add lines for labels. Keep your drawings neat!

FLORAL FORMULAS

Flowers are complex and there is enormous diversity in their structure. Floral formulas provide an easy way to
write down the important features of flowers and thereby capture structural diversity. Throughout the semester you
will need to interpret the floral formulas provided for each plant family as well as construct your own for the flower
specimens we will dissect in Lab. To construct a floral formula, always follow these steps:

1. Write the symbol for the flower’s gender. (HINT: Is


the flower perfect or imperfect?) Ca? Co? A?
♂ EPIGYNOUS
♀ HERMAPHRODITE Perfect Flowers G?
♂ STAMINATE Male Flowers
♀ CARPELLATE Female Flowers
Ca? Co? A?
EPIGYNOUS WITH
2. Identify the floral series present and how they are
arranged in relation to one another. G? HYPANTHIUM

Ca CALYX Sepals
Co COROLLA Petals 4. Find the superscript for each FLORAL SERIES by
A ANDROECIUM Stamen(s) counting the number of parts that make it up
G GYNOECIUM Carpel(s) (Hint: The subscripts can range from 0 (structure
not present) to ∞.) For example:

3. Determine the ovary INSERTION TYPE and use G 5 5 distinct carpels make up an
the appropriate format to construct your floral APOCARPOUS GYNOECIUM
formula. (Hint: Is the ovary superior or inferior
and is there a hypanthium?)
5. Determine if any of the FLORAL SERIES parts
HYPOGYNOUS are connate to one another. If so, draw a circle
Ca? Co? A? G? around their subscript. For example:

G ⑤ 5 connate carpels make up a


Ca Co A G
? ? ? ? PERIGYNOUS SYNCARPOUS GYNOECIUM

84 Integrative Biology 335: Plant Systematics


6. Determine if the flower’s Ca and Co have 7. If the flower has EPIPETALOUS STAMENS make
ACTINOMORPHIC or ZYGOMORPHIC symmetry. sure to indicate it in the floral formula as pictured
If actinomorphic symmetry, the floral formula below.
remains unchanged and if zygomorphic
symmetry add a Z next to Ca, Co or both. A?
EPIPETALOUS STAMENS
Ca Co A G
? ? ? ? Ca? Co? G?
ACTINOMORPHIC

CaZ? CoZ? A? G?
ZYGOMORPHIC

FLORAL MORPHOLOGY TERMINOLOGY LAB EXERCISE

The major objective of today's Lab exercise is to acquaint you with floral terminology. As in Lab 2: Vegetative
Terminology, there is a bewildering number of terms, but we will stress only the more commonly used ones in this
course. Make sure to learn these floral terms as soon as possible! The majority of the lab will be spent on plant
families and if you do not have the arsenal of floral vocabulary used to describe them, you will quickly become lost.

Begin today's Lab by reading over the Floral Formula examples on the next few pages. These will give you an idea
of how floral fomulas are constructed and interpreted. Afterward, complete the Floral Morphology Lab exercise by
examining, dissecting and answering questions about the flowers of three different plant species. Please use the
"Glossary of Floral Terminology" provided at the end of this Lab to help you.

FLORAL FORMULA EXAMPLES

Liriodendron tulipifera L. (Magnoliaceae) (Tulip Tree)

The flowers of L. tulipifera are perfect (thus plants are SYNOECIOUS) and complete
with ACTINOMORPHIC (RADIAL) floral symmetry. They have 3 distinct sepals in their
♂ Ca3 Co6 A∞ G∞
♀ calyx, 6 distinct petals in their corolla, numerous stamens in their androecium and
numerous carpels in their APOCARPOUS gynoecium. All floral series are free from
one another. The ovaries are superior and the floral insertion is HYPOGYNOUS.

Longitudinal Section (L. S.) of Flower

Lab 3 - Floral and Inflorescence Terminology 85


Geranium maculatum L. (Geraniaceae) (Spotted Geranium)

The flowers of G. maculatum are perfect (thus plants are SYNOECIOUS) and
complete with ACTINOMORPHIC (RADIAL) floral symmetry. They are considered
5-merous because they have 5 distinct green sepals in their calyx, 5 distinct pink
♂ Ca5 Co5 A10 G⑤
♀ petals in their corolla, 10 distinct stamens in their androecium and 5 connate
carpels in their SYNCARPOUS gynoecium. All floral series are free from one
another. The ovary is superior and the floral insertion is HYPOGYNOUS.

Citrullus lanatus (Thunb.) Matsum. & Nakai (Cucurbitaceae) (Watermelon)

The flowers of C. lanatus are imperfect and incomplete, because the STAMINATE
(male) flowers lack a gynoecium and the CARPELLATE (female) flowers lack
♂ Ca5 Co⑤ A⑤ G0 an androecium. Both genders of flower exhibit ACTINOMORPHIC (RADIAL) floral
symmetry and have 5 distinct sepals in their calyx and 5 connate petals in
their corolla. The 5 anthers of the STAMINATE flowers are connate (which is an
♀ Ca5 Co⑤ A0 unusual feature) and 3 connate carpels make up the SYNCARPOUS gynoecium
of CARPELLATE flowers. All floral series are free from one another. The ovary
G③ of the carpellate flowers is inferior and the floral insertion is EPIGYNOUS. Since
both male and female flowers are found on the same plant, the plants are
MONOECIOUS.

86 Integrative Biology 335: Plant Systematics


Asarum canadense L. (Aristolochiaceae) (Canadian Wild Ginger)

The flowers of A. canadense are perfect (thus plants are SYNOECIOUS) with
ACTINOMORPHIC (RADIAL) floral symmetry. However, these flowers are incomplete and
♂ Ca3 Co0 A12
♀ UNISERIATE because they have 3 distinct sepals in their calyx but have no corolla.
There are 12 distinct stamens in their androecium and 6 connate carpels in their
G⑥ SYNCARPOUS gynoecium. All floral series are free from one another. The ovary is
inferior and the floral insertion is EPIGYNOUS.

Cross Section (X. S.) of Ovary

Penstemon pallidus Small (Scrophulariaceae) (Pale Beardstongue)

These flowers are perfect (thus plants are SYNOECIOUS) and complete with
ZYGOMORPHIC (BILATERAL) floral symmetry. They have 5 connate sepals in their
calyx, 5 connate petals in their corolla, 4 stamens in their androecium and 2
A4 connate carpels in their SYNCARPOUS gynoecium. The ovary is superior and the
♂ CaZ⑤ CoZ⑤ G②
♀ floral insertion is HYPOGYNOUS. Also, the androecium is adnate to the corolla
(EPIPETALOUS STAMENS). This flower also has a staminode, which is not indicated
in the floral formula.

Lab 3 - Floral and Inflorescence Terminology 87


FLORAL MORPHOLOGY: SPECIMEN 1

Before you being examining any flowers, set up a


dissecting microscope and light source. Feel free to work
in pairs.

Crassula argentea Thunb. (CRASSULACEAE) (Jade plant)

Crassula is Latin and is the diminutive of "crassus,"


meaning thick in reference to the thick and fleshy (i.e.
succulent) leaves found on this plant. Although the
Crassulaceae family is found the world over, it has its
greatest diversity in South Africa. Many Crassulaceae
species are common in arid regions, usually in stony soil
or on rocky outcrops, hence why the family is often called
the Stonecrop family.

1. Observe your flower from the top. What is its floral


symmetry?

2. CALYX: Look at the base of the flower under your


dissecting scope and note the 5 small green lobes of
the sepals that comprise the calyx. Remember the
sepals alternate with the petals, meaning that the
sepals are found between the larger, white petals.

3. COROLLA: How many petals make up the corolla?


Note the small nectariferous appendage that is found
externally on each carpel near its base. What is the
function of these nectaries?

4. ANDROECIUM: Observe how the stamens are


alternating with the petals and are opposite the small
sepal lobes. How many stamens are there in the
androecium?

5. GYNOECIUM: The gynoecium of this flower is made


up of 5 distinct (un-fused) carpels. How many ovaries
are in the flower? Are there few or many ovules per
ovary?

GYNOECIUM TYPE ________________________________

OVARY POSITION ________________________________

INSERTION TYPE ________________________________

88 Integrative Biology 335: Plant Systematics


FURTHER QUESTIONS:

1. Dissect a carpel to determine placentation type. You will have to do this under your dissecting scope.

2. Based on your examination of the flower, is it COMPLETE or INCOMPLETE? PERFECT or IMPERFECT?

3. Below is a drawing of a longitudinal section (L.S.) of Crassula. Label all parts and construct a complete floral
formula for this flower. Also, try to verbally describe either in list or paragraph form your rational for your floral
formula. Make sure to use the appropriate terminology!

L. S. of Flower

FLORAL FORMULA

RATIONAL

Lab 3 - Floral and Inflorescence Terminology 89


FLORAL MORPHOLOGY: SPECIMEN 2

Nicotiana alata Link & Otto (SOLANACEAE)

Nicotiana is named for Jean Nicot (1530 - 1600), French


consul to Portugal, who is said to have first presented
tobacco to the courts of Portugal and France. Tobacco
is native to the Americas. The flower of Nicotiana usually
open at night and are then most fragrant. What might
pollinate this flower?

1. Observe your flower from the top. What is the floral


symmetry?

2. CALYX: Observe the pubescent calyx. In Nicotiana,


the calyx is often described as being toothed or cleft
(meaning deeply lobed). How many sepals make up
the calyx? Are they connate?

3. COROLLA: The corolla of this flower is described


as salverform because of its long, slender tube and
abruptly expanded flat limb. How many connate petals
make up the corolla tube? (Hint: Count the number of
corolla lobes.)

4. ANDROECIUM: How many EPIPETALOUS STAMENS


make up the androecium of this flower? Also, what
does the descriptive term EPIPETALOUS mean? Are the
stamens in anthesis (that is, is pollen being shed)?

5. GYNOECIUM: Note the bilobed stigma of the


gynoecium. How many carpels do you think fused to
produce this gynoecium? Make a cross section (X.S.)
of the ovary, draw it in Box 1 and label its parts. How
many locules do you see?
Box 1

GYNOECIUM TYPE ________________________________

OVARY POSITION ________________________________

INSERTION TYPE ________________________________

PLACENTATION ________________________________

90 Integrative Biology 335: Plant Systematics


FURTHER QUESTIONS:

1. Based on your observation of the flower, is it COMPLETE or INCOMPLETE? PERFECT or IMPERFECT?

2. Below is a drawing of a longitudinal section (L.S.) of Nicotiana. Label all parts and construct a complete floral
formula for this flower. Also, try to verbally describe either in list or paragraph form your rational for your floral
formula. Again, be sure to use the appropriate terminology!

L. S. of Flower

FLORAL FORMULA

RATIONAL

Lab 3 - Floral and Inflorescence Terminology 91


FLORAL MORPHOLOGY: SPECIMEN 3

Fuchsia triphylla L. (Onagraceae)

The family Onagraceae is cosmopolitan in distribution and


especially prominent in western North America. The family
includes many showy wildflowers and some ornamentals.
Fuchsia derives its name from the important physician and
botanist, Leonard Fuchs (1501 - 1565). Many physicians
of the Renaissance period and before were actually
practicing plant taxonomists. Why?

1. Observe the flower. What is the floral symmetry?

2. CALYX: In this species the calyx is petaloid (i.e. not


green and leafy) and its sepals are larger than the
petals. How many sepals make up the calyx?

3. COROLLA: Notice how the petals of the corolla


alternate with the sepals of the calyx. How many
petals make up the corolla? Collectively the calyx and
corolla make up the PERIANTH. Thus, the PERIANTH
possesses two whorls.

4. ANDROECIUM: Carefully cut the flower from top to


bottom using a sharp razor blade and examine the
stamens. How many are there? You’ll notice that they
are not all the same length. The stamens bearing the
shorter filaments are opposite the petals, while the
stamens bearing the longer filaments are opposite the
sepals.

The “floral tube” of this flower is more accurately


called a HYPANTHIUM. A HYPANTHIUM is a structure
derived by the adnation of the bases of the perianth
elements (calyx and corolla) and the androecium.
However, the sepals, petals and stamens are
separate from one another after they arise from the
rim of the hypanthium. At the base of the hypanthium
you should see four swollen glands or nectaries.
What is their purpose? (Hint: If there is a pool of liquid

92 Integrative Biology 335: Plant Systematics


remaining at the base of the hypanthium, have a
taste. It’s yummy.)

5. GYNOECIUM: The stigma of Fuchsia is swollen and


prominently 4-lobed. How many carpels make up the
gynoecium of the flower and what features did you
look at to determine carpel number?

GYNOECIUM TYPE ________________________________

OVARY POSITION ________________________________

INSERTION TYPE ________________________________

PLACENTATION ________________________________

FURTHER QUESTIONS:

1. Based on your observation of the flower, is it COMPLETE or INCOMPLETE? PERFECT or IMPERFECT?

2. Draw and label a longitudinal section (L.S.) of a F. triphylla flower as well as a cross section (X.S.) of its ovary
and construct its floral formula in the space provided.

FLORAL FORMULA

Lab 3 - Floral and Inflorescence Terminology 93


INFLORESCENCE TERMINOLOGY LAB EXERCISE

In this portion of today’s Lab exercise, you will become familiar with the parts, positions and types of various
INDETERMINATE and DETERMINATE inflorescences. Remember that an inflorescence is simply the arrangement of
flowers on a floral axis. If you’re unsure of a term, refer to the Inflorescence Type flow chart on the next page or the
"Glossary of Inflorescence Terminology" provided at the end of this Lab.

INFLORESCENCE PARTS

Just as leaves have an assortment of terms used to describe their


consitituent parts, so too do inflorescences. In the table to the INFLORESCENCE COMPOUND LEAF
right is a list of the parts of inflorescences and compound leaves
that often are mixed up by students (Too many terms that begin Flower Leaflet (Pinna)
with the letter P!). On the diagram below, please label these parts. Pedicel Petiolule
Also, label the BRACTS, BRACTEOLES and INVOLUCRE OF BRACTS
on the inflorescence. Rachis Rachis

Peduncle Petiole

INFLORESCENCE COMPOUND LEAF

94 Integrative Biology 335: Plant Systematics


INFLORESCENCE POSITION

Some plant species have characteristic inflorescence positions. For example, pineapples always display an
intercalary inflorescence position. Notice the vegetative material on the top of the fruit. Alternatively, redbud and
chocolate trees exhibit a cauliflory inflorescence position due to the placement of the flowers directly on the tree
branches. Make sure that you become familiar with all the inflorescence position terms found in the "Glossary of
Inflorescence Terminology"!

INFLORESCENCE TYPES

As with inflorescence positions, inflorescence types may be a key feature for a plant family. On display you will find
an assortment of live plants and/or herbarium specimens. Examine them and in the space provided below, record
the name of the specimen(s) that display(s) each of the major inflorescence types. Keep in mind that some of the
inflorescence types will only be represented by one specimen.

RACEME CAPITULUM

1. ___________________________________________ 1. ___________________________________________

2. ___________________________________________ 2. ___________________________________________

PANICLE SIMPLE UMBEL

1. ___________________________________________ 1. ___________________________________________

2. ___________________________________________ 2. ___________________________________________

SPIKE COMPOUND UMBEL

1. ___________________________________________ 1. ___________________________________________

2. ___________________________________________ 2. ___________________________________________

CORYMB CYME

1. ___________________________________________ 1. ___________________________________________

2. ___________________________________________ 2. ___________________________________________

Lab 3 - Floral and Inflorescence Terminology 95


96
INDETERMINATE DETERMINATE
OLDEST YOUNGEST
OLDEST YOUNGEST
FLOWER FLOWER FLOWER FLOWER

Progression of Flowering
Progression of Flowering

Progression of Flowering
Progression of Flowering
YOUNGEST OLDEST
YOUNGEST OLDEST
FLOWER FLOWER
FLOWER FLOWER

DETERMINATE INDETERMINATE
DETERMINATE INDETERMINATE

RACHIS ELONGATED RACHIS SHORTENED RACHIS FLATTENED RACHIS ABSENT

SIMPLE CORYMB
SIMPLE CORYMB COMPOUND CORYMB
COMPOUND CORYMB PANICLE PANICLE RACEME SPIKE
RACEME SPIKE

PEDICILLATE SESSILE
FLOWERS FLOWERS SIMPLE CYME COMPOUND CYME

HEAD (CAPITULUM) SIMPLE UMBEL COMPOUND UMBEL


SIMPLE CORYMB COMPOUND CORYMB PANICLE RACEME
HEAD (CAPITULUM) SPIKE SIMPLE UMBEL COMPOUND UMBEL

SIMPLE CORYMB COMPOUND CORYMB PANICLE ▪▪ ORRACEME SPIKE

COMPOUND CORYMB PANICLE RACEME SPIKE HEAD (CAPITULUM) SIMPLE UMBEL COMPOUND UMBEL

MPLE CORYMB
PANICLE COMPOUND CORYMB
RACEME SPIKE PANICLE RACEME SPIKE
throughout the semester.

HEAD (CAPITULUM) SIMPLE UMBEL COMPOUND UMBEL


INFLORESCENCE TYPES

Use this flow chart as a guide to


help identify the different types of
inflorescences you will encounter

Integrative Biology 335: Plant Systematics


SIMPLE UMBEL COMPOUND UMBEL
AD (CAPITULUM) COMPOUND SIMPLE
UMBEL UMBEL COMPOUND UMBEL
GLOSSARY OF FLORAL TERMINOLOGY

Listed below is the floral terminology that you will need to become skillful at using this semester. You should be able
to draw, compare and contrast each of these terms. You should also know the relationship between each of the terms
listed (i.e. CHARACTER STATES) and the major category (i.e. CHARACTER) to which they belong.

FLORAL SERIES COMPLETENESS

The four floral series terms listed below are COMPLETE


the collective terms for the major floral parts All four floral series present (Ca, Co, A & G)
(i.e. sepals, petals, stamens and carpels). For
example, all the sepals cumulatively make up INCOMPLETE
the flower’s CALYX). One or more floral series absent

PERFECT
1. CALYX Ca Sepals
Flower with both A and G functional
2. COROLLA Co Petals
3. ANDROECIUM A Stamen(s) IMPERFECT
4. GYNOECIUM G Carpel(s) Flower lacking either a functional A and G

STAMINATE
CALYX (SEPALS) Male flower; has a functional A but
lacks a functional G
COROLLA (PETALS)
CARPELLATE
Female flower; has a functional G but
ANDROECIUM (STAMENS) lacks a functional A

GYNOECIUM (CARPELS)

FLORAL DIAGRAM

ANTHER STIGMA
ANDROECIUM
(STAMENS) GYNOECIUM
STYLE
(CARPELS)
FILAMENT
OVARY

COROLLA (PETALS)

OVULE PERIANTH

CALYX (SEPALS)

PEDICEL RECEPTACLE
(or PEDUNCLE if solitary flower)

Lab 3 - Floral and Inflorescence Terminology 97


PLANT CONDITION FLORAL SYMMETRY

ACTINOMORPHIC (RADIAL)
SYNOECIOUS
Divisible into equal halves by two or more
All flowers on a plant are perfect
planes
MONOECIOUS
ZYGOMORPHIC (BILATERAL)
Both staminate and carpellate flowers occur on
Divisible into equal halves in one plane only
the same plant

DIOECIOUS
Staminate and carpellate flowers occur on
separate plants

ACTINOMORPHIC SYMMETRY ZYGOMORPHIC SYMMETRY

SYNOECIOUS MONOECIOUS DIOECIOUS


PERIANTH (CALYX AND COROLLA)

PERIANTH
Collective term used for the Ca and Co
FLORAL PART FUSION TERMINOLOGY
UNISERIATE
To determine the correct term to use to In one whorl or series (i.e. only Ca or only Co)
describe the fusion of floral parts, use the
chart below by answering the following two BISERIATE
questions: In two whorls or series (i.e. both Ca & Co)

CORONA
1. Do the floral parts belong to the same or An extra series of floral parts formed from
different floral series? outgrowths of the perianth parts, stamens or
receptacle. Often showy and diverse.
2. Are the floral parts fused to one another
or not? TEPALS
Term used to describe the sepals and petals
FLORAL PARTS when the two cannot be differentiated (e.g.
Same Different Cactaceae and Liliaceae)

3-MEROUS, 4-MEROUS, ETC.


Fused CONNATE ADNATE
FUSION

Indicates number of parts in a specific floral


series (e.g. a 4-MEROUS flower might have 4
Un-fused DISTINCT FREE
sepals, 4 petals, 8 stamens, and 4 carpels)

98 Integrative Biology 335: Plant Systematics


COROLLA TERMS ANDROECIUM

APETALOUS STAMENS
Lacking petals The male reproductive unit (the site of pollen
production). One to many separate or fused
SYMPETALOUS STAMENS comprise an ANDROECIUM.
Petals that are connate at least at the base
FILAMENT
BILABIATE COROLLA Stalk of a stamen
Corolla is two-lipped (e.g. Lamiaceae and
(sometimes) Scrophulariaceae) ANTHER
Pollen producing portion of a stamen
PLICATE COROLLA (meiosis occurs within the pollen sacs)
Corolla is folded like a fan (e.g. Solanaceae)
POLLEN
The male gametophytes

CONNECTIVE
The sterile tissue connecting the two
locules of an anther

CONNIVENT ANTHERS
Anthers that converge but that are not actually
connate to one other (e.g. Solanaceae)
BILABIATE COROLLA PLICATE COROLLA

VERSATILE ANTHERS
Anthers attached to filaments in their center
rather than at one of their ends (e.g. Liliaceae)
COROLLA SHAPE VISCIN THREADS
A clear, sticky substance that holds together
CAMPANULATE large numbers of pollen grains (e.g.
Bell-shaped (e.g. Ericaceae) Onagraceae)

URCEOLATE
Urn or pitcher-shaped (e.g. Ericaceae)
CONNECTIVE
ANTHER

FILAMENT

STAMEN FRONT VIEW STAMEN BACK VIEW

CAMPANULATE URCEOLATE ANDROECIUM

Lab 3 - Floral and Inflorescence Terminology 99


SPECIAL TYPES OF STAMENS

LAMINAR STAMENS MONADELPHOUS STAMENS


Ancestral stamens that possess wide and Stamen filaments connate to form one staminal
flattened ANTHERS and very tiny FILAMENTS (e.g. tube (e.g. Malvaceae)
Magnoliaceae)
DIADELPHOUS STAMENS
EPIPETALOUS STAMENS Stamens united into two, often unequal, sets by
Stamens that are fused (adnate) to corolla. Key selective connation among their filaments (e.g.
feature of the families in the Asterid clade. Fabaceae subfamily Faboideae (9 + 1))

DIDYNAMOUS STAMENS STAMINODES


Having two short and two long stamens (e.g. Sterile stamens (they do not produce pollen);
Lamiaceae and Scrophulariaceae) variable in form and size

TETRADYNAMOUS STAMENS STAMINODIUM


Having two short and four long stamens (e.g. The 5th sterile stamen in Scrophulariaceae
Brassicaceae)

LAMINAR STAMENS EPIPETALOUS STAMENS TETRADYNAMOUS STAMENS (4 LONG + 2 SHORT)

MONADELPHOUS STAMENS DIADELPHOUS STAMENS DIDYNAMOUS STAMENS


(9 CONNATE STAMENS + 1 DISTINCT STAMEN) (2 LONG + 2 SHORT)

100 Integrative Biology 335: Plant Systematics


GYNOECIUM PARTS GYNOECIUM TYPES

CARPELS MONOCARPOUS GYNOECIUM


The female reproductive unit (the site of ovule One carpel makes up the gynoecium
production, pollination and fertilization). One
to many separate or fused carpels comprise a APOCARPOUS GYNOECIUM
gynoecium. Multiple (>1) distinct carpels make up the
gynoecium
STIGMA
The pollen receptive portion of the gynoecium. SYNCARPOUS GYNOECIUM
Multiple (>1) connate carpels make up the
STYLE gynoecium
The elongated portion between stigma and
ovary specialized for pollen tube growth.

OVARY
Basal portion that surrounds and protects the
ovules.

LOCULE
The chamber(s) within an ovary

SEPTUM (SEPTA)
Interior wall which separates the locules in
instances where 2 or more chambers occur
(walls = SEPTA)

PLACENTA (pl. PLACENTAE)


The region or line along which the ovules are
MONOCARPOUS APOCARPOUS SYNCARPOUS
attached

OVULE
Structure containing the female gametophyte
(meiosis occurs within the ovule). Matures into
a seed OVARY POSITION AND INSERTION TYPE

FUNICULUS SUPERIOR OVARY


The stalk connecting the ovule to the placenta Ovary situated above the point of attachment
of the perianth and androecium and wholly free
from them (stamens may be adnate to corolla)
STIGMA
HYPOGYNOUS
A flower with perianth and androecium
STYLE arising from below the ovary (gynoecium)

PERIGYNOUS
A flower with perianth and androecium
arising from a HYPANTHIUM (floral cup) that
OVULE is not adnate to the ovary. (Remember the
FUNICULUS ovary is still superior!)
OVARY
PLACENTA INFERIOR OVARY
LOCULE Ovary is below the point of attachment of the
perianth and androecium. In other words, the
SEPTUM outer floral whorls are adnate to the ovary. A
HYPANTHIUM (floral cup) may or may not be
present.
GYNOECIUM

Lab 3 - Floral and Inflorescence Terminology 101


OVARY POSITION & INSERTION TYPE (cont.) PLACENTATION TYPE (cont.)

EPIGYNOUS In SYNCARPOUS gynoecia:


A flower with perianth and androecium
arising upon the ovary rather than the AXILE
receptacle. The placental area of the ovary is attached
to an axis derived from the connate
margins of the component carpels - such
an ovary is divided into two or more
locules by septa. The ovules are borne
along the central axis.

PARIETAL
The placental areas are attached to the
side walls of the ovary (or extrusions of
the wall) - such an ovary usually has one
HYPOGYNOUS PERIGYNOUS
locule (therefore no septa).

APICAL
Attachment of ovules to the top of the
ovary (one locule - no septa).

BASAL
Attachment of ovules to the bottom of the
ovary (one locule - no septa).

FREE-CENTRAL
Attachment of ovules to a free-standing
EPIGYNOUS EPIGYNOUS (WITH HYPANTHIUM) central column in the ovary (one locule -
no septa).

PLACENTATION TYPE

PLACENTATION
The arrangement of ovules within the ovary. AXILE PARIETAL
To determine the type of placentation, first
determine the gynoecium type (certain
placentation types are found in certain
gynoecium types).

In MONOCARPOUS or APOCARPOUS gynoecia:

MARGINAL
APICAL BASAL
The ovules are attached to the folded
margins of the carpel.

FREE-CENTRAL
MARGINAL

102 Integrative Biology 335: Plant Systematics


MISCELLANEOUS TERMS

ANDROGYNOPHORE (i.e. GYNOPHORE)


Stalk that supports the corolla, androecium and
gynoecium (e.g. Caryophyllaceae)

EPICALYX
A whorl of bracts that subtends the sepals (e.g.
Malvaceae)

GYNOSTEGIUM
A structure formed from the adnation of the
anthers of the androecium and stigmas of the
gynoecium (e.g. Asclepiadaceae)

HYPANTHIUM
A floral cup formed from the adnation of the
calyx, corolla and androecium

INVOLUCRE
A whorl of bracts subtending a flower or flower
cluster (e.g. Fagaceae)

ANDROGYNOPHORE

EPICALYX

HYPANTHIUM

Lab 3 - Floral and Inflorescence Terminology 103


GLOSSARY OF INFLORESCENCE TERMINOLOGY

Listed below is the inflorescence terminology that you will need to become skillful at using this semester. You should
be able to draw, compare and contrast each of these terms. You should also know the relationship between each of
the terms listed (i.e. CHARACTER STATES) and the major category (i.e. CHARACTER) to which they belong.

INFLORESCENCE PARTS

BRACTS FLOWER

A modified or much-reduced leaf associated


with an inflorescence or flower. These may BRACT
differ substantially from foliage leaves. RACHIS

BRACTEOLES or BRACTLETS PEDICEL


A smaller BRACT (or BRACTLET)
BRACTEOLES

INVOLUCRE
A series of BRACTS immediately subtending a
flower or inflorescence

PEDICEL
The stalk of one flower in an inflorescence of
INVOLUCRE
multiple flowers OF BRACTS

PEDUNCLE
The stalk of an inflorescence or a solitary
flower
PEDUNCLE
RACHIS
The primary axis of an elongated inflorescence

SEQUENCE OF FLOWERING

DETERMINATE INFLORESCENCE OLDEST YOUNGEST


An inflorescence in which the terminal or FLOWER FLOWER

central flower opens first, resulting in the


cessation of primary axis elongation.
Progression of Flowering

Progression of Flowering

INDETERMINATE INFLORESCENCE
An inflorescence in which the lowermost or
outermost flowers open first, with the primary
axis often elongating as the flowers develop.
Usually no terminal flower is produced.

YOUNGEST OLDEST
FLOWER FLOWER

DETERMINATE INDETERMINATE

104 Integrative Biology 335: Plant Systematics


DETERMINATE INFLORESCENCES

SIMPLE CYME
A three-flowered cluster composed of a
peduncle bearing a terminal flower and, below
it, two bracts with each bract subtending a
lateral flower.

COMPOUND CYME
A branched simple cyme.
SIMPLE CYME COMPOUND CYME

INDETERMINATE INFLORESCENCES

CORYMB RACEME
A relatively flat topped inflorescence with the lower Stalked flowers arranged along an elongate
pedicels longer than the upper. central axis (rachis).

HEAD (CAPITULUM) SPIKE


A compact inflorescence composed of a very short Sessile flowers arranged along an elongate central
axis and usually sessile flowers. axis (rachis).

PANICLE UMBEL
Similar to a raceme but greatly branched. Several branches radiating from the same point
and are terminated by single flowers (Simple
umbels) or 2° umbels (compound umbels).

SIMPLE CORYMB COMPOUND CORYMB PANICLE RACEME SPIKE

HEAD (CAPITULUM) SIMPLE UMBEL COMPOUND UMBEL

Lab 3 - Floral and Inflorescence Terminology 105


INFLORESCENCE POSITION

AXILLARY INTERCALARY
Inflorescence arises from the axil of a leaf or bract Inflorescence seemingly in the middle of a
stem. Essentially, the main stem axis continues
BASAL to grow vegetatively after having produced an
Inflorescence where flowers are borne on a inflorescence
leafless peduncle that arises from a cluster of
leaves at the base of the plant TERMINAL
Inflorescence terminates a stem
CAULIFLORY
Flowers that grow directly upon woody stems or WHORLED
trunks Inflorescence where there are multiple flowers
arranged in a circle (whorl) around a single node
(verticil)

AXILLARY BASAL

TERMINAL

CAULIFLORY

INTERCALARY WHORLED (VERTICIL)

106 Integrative Biology 335: Plant Systematics


SPECIAL INFLORESCENCE TYPES

CATKIN (AMENT) SPADIX


An inflorescence consisting of a dense spike A spike with small flowers crowded on a thickened
or raceme of apetalous, unisexual flowers (e.g. axis that is subtended by a SPATHE (e.g. Araceae)
Betulaceae, Fagaceae, Salicaceae).
SPIKELET
CYATHIUM (pl. CYATHIA) The inflorescence of grasses and sedges
The inflorescence of the genus Euphorbia in consisting of one to many flowers subtended by
Euphorbiaceae, consisting of a cup-like involucre two bracts (glumes) (e.g. Poaceae).
containing a single female flower (just gynoecium,
no calyx, corolla or androecium) and male flowers SYCONIUM (pl. SYCONIA)
with a single stamen. Fig inflorescence consisting of a hollow, inverted
receptacle containing simple, unisexual flowers.

FEMALE
FLOWER
SPADIX

MALE SPATHE
FLOWER

CATKIN

SPADIX

CYATHIUM

FLORET

MALE FLOWERS
PALEA

AWN FEMALE FLOWERS

FLESHY SEPALS OF
LODICULES FEMALE FLOWERS

LEMMA
RACHILLA

2° GLUME 1° GLUME
(UPPER GLUME) (LOWER GLUME)

PEDUNCLE SYCONIUM

SPIKELET

Lab 3 - Floral and Inflorescence Terminology 107


108 Integrative Biology 335: Plant Systematics
LABORATORY 4
FRUIT TERMINOLOGY

FRUIT AND SEED MORPHOLOGY

▪▪ A FRUIT is a ripened (mature) ovary of a flower along with any adnate parts.
▪▪ A SEED is a ripened (or fertilized) ovule containing an embryo within a seed coat and often with additional storage tissue.

The term “fruit” originally meant “any plant used as food”. Since the ancient Greeks, fruits have been eaten at the
end of the meal and not with the meal. Consequently, most people commonly view fruits as sweet, soft plant products
used in dessert foods. Alternatively, vegetables (the leaves, tubers, roots or even entire inflorescences of plants) are
considered savory or main course foods. Though it is true that fruits generally have a much higher sugar content and
are often more acidic those those things we call vegetables, botanically speaking not all fruits are soft and sweet and
many are not good to eat at all!

A more precise, technical definion for a fruit was established as the science of botany arose in the 18th century.
Botanically, a fruit is a mature ovary together with any floral or vegetative structures that are attached to it and that
become enlarged and ripen with it. Additionally, the wall of the ovary becomes the PERICARP (the wall of the fruit).
The OVULES become the SEEDS. The ZYGOTE matures into the EMBRYO, the INTEGUMENTS mature into the SEED COAT,
and the FUNICULUS develops into the SEED STALK.

Taxonomists have developed a usable classification


FLOWER PART FRUIT PART of fruits based primarily on features such as texture,
▪▪ Ovary wall ▪▪ Pericarp
nature of dehiscence, number of seeds, presence
or absence of attached parts, and the morphology
▪▪ Ovule ▪▪ Seed of the ovary from which the fruits were derived.
Today in Lab you will learn the major fruit types
▪▪ Integuments (outer covering of ovule) ▪▪ Seed coat based on morphology and taxonomy as well as
▪▪ Zygote ▪▪ Embryo
begin to examine the characteristics of the flowers
they arise from.
▪▪ Fusion Nucleus ▪▪ Endosperm

▪▪ Funiculus (stalk of an ovule) ▪▪ Seed Stalk

MORPHOLOGY-BASED FRUIT TYPES

When classifying fruits based on morphology, there are three main groups:

1. DRY, DEHISCENT FRUITS


2. DRY, INDEHISCENT FRUITS
3. FLESHY FRUITS

Lab 4 - Fruit Terminology 109


110
DRY FLESHY

Pericarp splits open [seed Pericarp remains closed


is dispersal unit] [fruit is dispersal unit] BERRY DRUPE
Usually syncarpous G Superior ovary
Superior or inferior Usually contains 1 seed
DEHISCENT FRUIT INDEHISCENT FRUIT ovary
Has distinctive exo-,
Entire pericarp fleshy meso- and endocarp

FOLLICLE CAPSULE SCHIZOCARP ACHENE NUT


Apocarpous G Syncarpous G Syncarpous G Variable G Syncarpous G
[from one carpel]
Contains few to Splits into 2 or more 1-seeded with 1-seeded with a
Opens along a many seeds 1-seeded segments seed free from thick, bony
single suture [mericarps] pericarp pericarp
The most common
and variable of all Involucre present
fruit types

NUTLET
A small nut without
the involucre
their morphology.

DEHISCENCE TYPE SAMARA


A winged achene

SEPTICIDAL LOCULICIDAL PORICIDAL DENTICIDAL CIRCUMSCISSLE

Splits along septa Splits along locule Seeds released Seeds released Top comes off [like
through pores through apical lid] to release seeds
MORPHOLOGY-BASED FRUITS

teeth
you will encounter throughout the semester based on
Use this flow chart as a guide to help identify the fruits

Integrative Biology 335: Plant Systematics


DRY AND DEHISCENT FRUITS

FOLLICLE

▪▪ Asclepias incarnata L. (Swamp Milkweed)


▪▪ Asclepias syriaca L. (Common Milkweed)
▪▪ Aquilegia spp. (Columbines)
▪▪ Spiraea spp. (Meadowsweet)

There are many plant species that produce FOLLICLE fruits


including: milkweed, larkspur, columbine and peony. Observe the
specimens on display. Notice how FOLLICLES split along one side
only, exposing the seeds within. FOLLICLES are derived from one
carpel. Based on this, what type or types of gynoecium could a
flower have to produce this fruit type?

CAPSULE

▪▪ Aristolochia spp. (Dutchman’s Pipe)


▪▪ Datura stramonium L. (Datura)
▪▪ Glaucium spp. (Horned Poppy)
▪▪ Hamamelis mollis Oliv. (Witch Hazel)
▪▪ Hemerocallis spp. (Day Lily)
▪▪ Hibiscus spp. (Hibiscus)
▪▪ Oenothera speciosa Nutt. (Pinkladies)
▪▪ Papaver spp. (Poppies)
▪▪ Portulaca oleracea L. (Common Purslane)
▪▪ Silene spp. (Catchfly or Campion)
▪▪ Yucca spp. (Yucca)

CAPSULES are the most common of the dried fruit types that
dehisce. They are derived from a syncarpous gynoecium, are
typically several to many-seeded, consist of at least 2 carpels and
split in a variety of ways.

Based on their mode of dehiscence (how they release seed),


capsules have been classified into five different types. SEPTICIDAL
CAPSULES split along the partitions or septa between their carpels,
while LOCULICIDAL CAPSULES split open through the locules to
dehisce. PORICIDAL CAPSULES have a row of pores through which
the seed are shaken out as the fruits rattles in the wind. DENTICIDAL
CAPSULES release their seeds through apical teeth. CIRCUMSCISSLE
CAPSULES (or PYXIS) form a cap towards one end that pops off
during dehiscence.

Lab 4 - Fruit Terminology 111


For each capsule type below, list the names of the fruits on
display that exhibit that type of dehiscence.

SEPTICIDAL CAPSULE

__________________________________________________

__________________________________________________

__________________________________________________

LOCULICIDAL CAPSULE

__________________________________________________

__________________________________________________

__________________________________________________

PORICIDAL CAPSULE

__________________________________________________

__________________________________________________

__________________________________________________

DENTICIDAL CAPSULE

__________________________________________________

__________________________________________________

__________________________________________________

CIRCUMSCISSLE CAPSULE

__________________________________________________

__________________________________________________

__________________________________________________

Differentiating between loculicidal and septicidal capsules can


Box 1 sometimes be quite tricky. However, if you think about the fruits
in terms of their cross sections, it will be much easier. In Box 1,
sketch a cross section of a loculicidal capsule and a septicidal
capsule. Make sure these drawings clearly show where the
fruit splits and where the fruit’s septa are located.

112 Integrative Biology 335: Plant Systematics


DRY AND INDEHISCENT FRUITS

ACHENE

▪▪ Fagopyrum esculentum Moench (Buckwheat)


▪▪ Helianthus spp. (Sunflower)
▪▪ Polygonum spp. (Smartweed)

An ACHENE is a dry, indehiscent fruit type that is extremely variable


in form and is derived from a monocarpous, apocarpous or
syncarpous gynoecium. It is composed of a single seed attached
only at its base to the fruit’s hard PERICARP; therefore the seed is
very easily removed from its pericarp. What is a PERICARP and what
did it arise from in the original flower?

Crack open a sunflower seed or observe one on display. What are


you actually cracking open?

NUT

▪▪ Castanea spp. (Chestnuts)


▪▪ Corylus americana Marshall (American Hazelnut)
▪▪ Quercus spp. (Oaks)

NUTS are dry, indehiscent fruits derived from a syncarpous


gynoecium, but are one-seeded by the abortion of ovules. They are
similar to ACHENES, but are generally larger with a much harder and
thicker PERICARP. Additionally, nuts mature with a cup (INVOLUCRE)
or cluster of bracts at their base. This semester you’ll learn about
two plant families that produce NUTS, Fagaceae (the Beech family)
and Betulaceae (the Birch family).

Many nuts in the popular sense are not nuts, botanically speaking.
Which of the following are true nuts?

Peanuts, coconuts, almonds, walnuts, pecans, brazil nuts,


cashews, pistachios

Answer: None are true nuts. A peanut is an atypical legume,


coconuts and almonds are drupes, and walnuts and pecans are
also drupes whose “flesh” withers and dries after the seed matures.
Brazil nuts are the seeds of a large capsule, and a cashew is the
single seed of a particular drupe. Pistachios are also the seeds of
drupes.

Lab 4 - Fruit Terminology 113


SCHIZOCARPS

▪▪ Anethum graveolens L. (Dill)


▪▪ Apium graveolens L. (Celery)
▪▪ Cuminum cyminum L. (Cumin)
▪▪ Foeniculum vulgare Mill. (Fennel)

SCHIZOCARPS are dry, indehiscent fruits that break into one-


carpellate segments called MERICARPS. Each MERICARP can contain
one to a few seeds, and the seeds remain completely surrounded
by the PERICARP. What type of gynoecium do SCHIZOCARPS develop
from?

Members of the carrot family, Apiaceae, are characterized by


this type of fruit. However, some species in Malvaceae produce
SCHIZOCARPS as well, though their morphology is somewhat
different. SCHIZOCARPS can also be winged as in Aceraceae (the
maple family).

SAMARA

▪▪ Acer spp. (Maples)


▪▪ Alnus glutinosa (L.) Gaertner (Black Alder)
▪▪ Fraxinus pennsylvanica Marshall (Green Ash)

In SAMARAS, the PERICARP around the seed extends out into a wing
or membrane. Essentially, a SAMARA is a winged ACHENE. What
function does the wing serve?

Note: SAMARAS can be produced singly as in ashes, elms and


birches or in pairs as in maples. For maples, a more precise term
for the paired fruit is a SAMAROID SCHIZOCARP (and not, as we’ve
seen written a samurai schizocarp) since the two samaras (or one-
carpellate segments (MERICARPS)) break apart.

114 Integrative Biology 335: Plant Systematics


FLESHY FRUITS

BERRY

▪▪ Persea americana Mill (Avocado)


▪▪ Musa acuminata Colla (Banana)
▪▪ Capsicum annuum L. (Bell Pepper)
▪▪ Vaccinium corymbosum L. (N. Highbush Blueberry)
▪▪ Solanum melongena L. (Eggplant)
▪▪ Vitis vinifera L. (Common Grape)
▪▪ Actinidia deliciosa C.F. Liang & A.R. Ferguson (Kiwi)
▪▪ Carica papaya Marsh (Papaya)
▪▪ Physalis philadelphica Lam. (Tomatillo)
▪▪ Lycopersicon esculentum Miller. (Tomato)

BERRIES typically develop from syncarpous gynoecia. Their entire


pericarp is fleshy and relatively soft at maturity. Although most
BERRIES contain more than one seed, notable exceptions are dates
and avocados, which have only one.

Berries can arise from flowers with superior or inferior ovaries. How
can you tell whether a berry has arisen from a superior or inferior
ovary?

Which of the fruits on display were derived from flowers with


superior ovaries?

Box 2
Which ones were derived from flowers with inferior ovaries?

Make or observe a cross section (X.S.) through a cherry tomato


and draw it in Box 2. What is the placentation type? How many
carpels fused to produce the gynoecium of a tomato flower? How
do you know?

What is the origin of the green, papery structure that surrounds a


tomatillo fruit?

Lab 4 - Fruit Terminology 115


DRUPE

▪▪ Cocos nucifera L. (Coconut)


▪▪ Olea europaea L. (Olive)
▪▪ Prunus americana Marsh (American Plum)
▪▪ Prunus armeniaca L. (Apricot)
▪▪ Prunus avium (L.) L. (Sweet Cherry)
▪▪ Prunus dulcis (Mill.) D.A. Webb (Sweet Almond)
▪▪ Prunus persica (L.) Batsch (Peach/Nectarine)

DRUPES usually develop from flowers with a superior ovary


containing a single ovule. Unlike in BERRIES, the PERICARP of
Box 3 DRUPES is differentiated into an EXOCARP, MESOCARP, and stony
ENDOCARP (the pit). Examine the cut fruit on display. Sketch one
of them out in Box 3, and then identify and label the following
structures: EXOCARP, MESOCARP, ENDOCARP and SEED.

The MESOCARP of a drupe is not always obviously fleshy. For


example, in coconuts the husk (consisting of the MESOCARP and
EXOCARP), is very fibrous and is usually removed before the rest
of the fruit is sold in markets. What purpose does this fibrous
MESOCARP serve? The edible portion of a coconut is the seed
and comprises both solid “meat” and liquid “milk” endosperm. The
endosperm is surrounded by the thick, hard ENDOCARP typical of
drupes.

Almonds are typically sold shelled, that is after the shells (or
endocarps) are removed. Blanched almonds are shelled almonds
that have been treated with hot water to soften and remove the
seedcoat.

Olives are a naturally bitter fruit that need to be fermented or cured


with lye or brine to make them palatable.

116 Integrative Biology 335: Plant Systematics


FRUIT TYPES BASED ON TAXONOMY

CARYOPSIS

▪▪ Hordeum vulgare L. (Common Barley)


▪▪ Secale cereale L. (Cereal Rye)
▪▪ Triticum aestivum L. (Common Wheat)
▪▪ Zea mays L. (Corn)

CARYOPSIS fruits are restricted to Poaceae (the Grass Family). They


are dry, indehiscent and very similar to ACHENES except that the
seed coat is fully adnate to the ovary wall (PERICARP). Think of a
corn kernel and how difficult it would be to separate the outer part
of the kernel from the inner! To review, what does adnate mean?

CYPSELA (CYPSELAE)

▪▪ Helianthus spp. (Sunflower)


▪▪ Liatris spp. (Blazing Star)
▪▪ Taraxacum officinale F.H. Wigg (Common Dandelion)
▪▪ Tragopogon spp. (Goat’s Beard)

CYPSELA fruits are restricted to Asteraceae (the Sunflower Family).


They are also dry, indehiscent and very similar to ACHENES except
that they usually have an adnate calyx. In Liatris, the calyx is
modified into bristle-like appendages called the PAPPUS. Are
CYPSELA fruits formed from a superior or inferior ovary?

Note: A sunflower fruit (commonly referred to as a sunflower seed)


is correctly termed a CYPSELA but does not possess an adnate
calyx. Consequently, it can be referred to as an ACHENE.

HESPERIDIUM

▪▪ Citrus japonica Thunb. (Kumquat)


▪▪ Citrus × limon (L.) Burm. (Lemon)
▪▪ Citrus × paradisi Macfad. (Grapefruit)
▪▪ Citrus × sinensis (L.) Osbeck (Sweet Orange)
▪▪ Citrus spp. (Lime)

HESPERIDIUM fruits are produced by most species in Rutaceae (the

Lab 4 - Fruit Terminology 117


Citrus Family). They are fleshy, indehiscent and very similar to
BERRIES except that they have a leathery rind containing oil glands
and numerous septa. Each segment of a HESPERIDIUM represents
a single carpel from the original flower’s gynoecium. Inside these
segments are fleshy hairs (outgrowths from the inner lining of the
ovary wall) that became swollen with water as the HESPERIDIUM
developed.

Examine the cut orange on display. Is it derived from a superior or


inferior ovary? How many carpels matured to produce it? What is
its placentation type?

LEGUME

▪▪ Arachis hypogaea L. (Peanut)


▪▪ Cercis canadensis L. (Eastern Redbud)
▪▪ Gleditsia triacanthos L. (Honey Locust)
▪▪ Gymnocladus dioicus (L.) K. Koch (Kentucky Coffeetree)
▪▪ Phaseolus vulgaris L. (Common Bean)
▪▪ Pisum sativum var. saccharatum L. (Snow Pea)

LEGUME fruits are produced by literally thousands of species in


Fabaceae (the Legume Family). They are dry, dehiscent and very
similar to FOLLICLES except that they usually split along two sutures
(seams) rather than one. What type of gynoecium is required to
produce a LEGUME? What is a LEGUME’s placentation type?

Dissect a bean or snow pea. Sketch it out in Box 4, and then


identify and label the following structures: SEEDS, FUNICULUS,
SUTURES and PERICARP.

Box 4

118 Integrative Biology 335: Plant Systematics


Peanuts are also LEGUMES, but they are atypical in that fruits
Box 5
develop and mature underground; the seeds are usually released in
nature by bacterial breakdown of the PERICARP instead of through
an active splitting action.

Crack open a peanut, sketch it in Box 5 and label the following:


PERICARP, seed(s), TESTA or SEED COAT, COTYLEDONS, PLUMULE
and EMBRYO. You’ll notice that most of the seed’s volume is taken
up by the cotyledons, two structures modified to store the energy
and protein necessary to get the seedling established. Where
is the endosperm? In the peanut (and other legume fruits), the
endosperm is absorbed by the embryo, which repackages the
nutrients in the cotyledons.

Fun Facts: The high fat content in peanuts renders them especially
vulnerable to rancidity and the acquisition of odors from their
surroundings. To prevent this, peanuts are roasted. Have you ever
eaten a raw peanut? For your general information, the peanut
is a native of South America, and has been found in Peruvian
settlements dated at 800 B.C.E.

PEPO

▪▪ Citrullus lanatus (Thunb.) Matsum. & Nakai (Watermelon)


▪▪ Cucurbita moschata Duchesne. (Butternut Squash)
▪▪ Cucumis sativus L. (Garden Cucumber)
▪▪ Cucumis melo subsp. melo (Canteloupe)
▪▪ Cucurbita pepo L. (Field Pumpkin)

PEPO fruits are restricted to Cucurbitaceae (the Cucumber Family).


They are fleshy, indehiscent and considered to be a special type of
BERRY due to their tough outer rind. Examine the fruit on display,
identify the following structures and label them on the cucumber
cross section below: SEEDS, PLACENTA and PERICARP. Are PEPOS
derived from inferior or superior ovaries? How can you tell?

Observe the sectioned cucumber on display. This fruit arose from


a syncarpous gynoecium of how many connate carpels? What
placentation type, characteristic of a PEPO, does the cucumber
display?

Lab 4 - Fruit Terminology 119


POME

▪▪ Cydonia oblonga Mill. (Quince)


▪▪ Malus pumila Mill. (Paradise Apple)
▪▪ Pyrus communis L. (Common Pear)

POME fruits are restricted to Rosaceae Subfamily Maloideae (the


Apple Subfamily). The bulk of the flesh of an apple or pear comes
from the enlarged fleshy hypanthium that grows up and around
the ovary. In the prepared cross-section of an apple, identify the
transition between the regions derived from the ovary (papery
tissue) and the hypanthium.

Are POMES derived from a superior or inferior ovary? Explain your


reasoning. Also, why is a POME also considered an ACCESSORY
FRUIT?

SILIQUE AND SILICLE

▪▪ Arabidopsis thaliana (L.) Heynh. (Mouse Ear Cress)


▪▪ Hesperis matronalis L. (Dame’s Rocket)
▪▪ Lunaria spp. (Money Plant)
▪▪ Thlaspi arvense L. (Field Pennycress)

SILICLE and SILIQUE fruits are restricted to Brassicaceae (the


Mustard Family). Superficially, these dry, dehiscent fruits may
look like FOLLICLES or LEGUMES, but possess a number of notable
characteristics that set them apart. Unlike FOLLICLES and LEGUMES,
SILICLES and SILIQUES arise from a syncarpous gynoecium. If these
fruits possess two locules, how many connate carpels comprise the
gynoecium?

Also, though SILICLES and SILIQUES split along two sutures (sides)
like LEGUMES, the seeds are borne on a central partition called
the REPLUM, which is exposed when the two halves of the fruit
separate. Observe the Lunaria specimen on display - only the
REPLUMS are left! In this case the valves have already fallen off,
thereby releasing the seeds.

Based on your observation of the display fruits, what do you think is


the key difference between a SILIQUE and a SILICLE?

120 Integrative Biology 335: Plant Systematics


SPECIAL FRUIT TYPES

ACCESSORY FRUIT

▪▪ Fragaria × ananassa (Strawberry hybrid)


▪▪ Nelumbo lutea Willd. (American Lotus)

ACCESSORY FRUITS possess tissue in addition to the ovary. In many


cases, such as Nelumbo and strawberries, the receptacle is the
tissue that becomes enlarged.

Examine the ACCESSORY FRUIT of Nelumbo, an aquatic plant


commonly called Lotus. Its flowers are long, pedunculate, perfect
and possess 20-30 tepals and numerous stamens (200 - 400).
What does pedunculate mean and what is a tepal?

The gynoecium of a Nelumbo flower consists of many distinct


carpels that are individually sunken in (but free from) the enlarged,
spongy receptacle. The ACHENE-like fruits within the persistent
receptacle have a small respiratory pore at the top. What’s the
purpose of this enlarged receptacle? How do you think this fruit is
dispersed?

In a strawberry, the cone-shaped receptacle becomes fleshy


and red, while the mature ovary of each carpel becomes a little
achene on its surface. Have a close look at the strawberry under
your dissecting scope. Identify the achenes on the surface of the
strawberry. Can you see any styles? Where are they?

Where are (were) the stamens, sepals and petals?

Why have strawberries elaborated such a mechanism for seed


dispersal?

Lab 4 - Fruit Terminology 121


AGGREGATE FRUIT

▪▪ Fragaria × ananassa (Strawberry hybrid)


▪▪ Rubus idaeus L. (American Red Raspberry)
▪▪ Liriodendron tulipifera L. (Tulip Tree)
▪▪ Magnolia grandiflora L. (Magnolia)

AGGREGATE FRUITS are derived from a single flower with an


apocarpous gynoecium of few to many distinct carpels. The
individual carpels each develop into tiny fruits or “fruitlets” (e.g.
drupelets), but they mature as a clustered unit on a single
receptacle.

Can a strawberry also be an AGGREGATE FRUIT? Why?

Examine the AGGREGATE FRUIT of Magnolia. Can you identify


the FOLLICLES? What evidence do you have suggesting this fruit
matured from only one flower?

Magnolia produces an AGGREGATE OF FOLLICLES; a raspberry an


AGGREGATE OF DRUPELETS (small drupes), Liriodendron produces
an AGGREGATE OF SAMARAS. How are these fruits dispersed?

MULTIPLE FRUIT

▪▪ Ananas comosus (L.) Merr. (Pineapple)


▪▪ Liquidambar styraciflua L. (Sweetgum)

MULTIPLE FRUITS arise from the maturation of individual ovaries from


many flowers along with accessory tissue (i.e the inflorescence
axis, calyx, and receptacle). Identify bracts and persistent sepals
on the pineapple. What evidence do you see indicating that the
pineapple developed from an intercalary inflorescence position?

Sweetgum is a MULTIPLE FRUIT OF LOCULICIDAL CAPSULES. The


seeds have most likely already been released from the specimens
on display.

122 Integrative Biology 335: Plant Systematics


GLOSSARY OF FRUIT TERMINOLOGY

Listed below is the fruit terminology that you will need to become skillful at using this semester. You should be able to
draw, compare and contrast each of these terms.

MORPHOLOGY-BASED: FLESHY FRUITS

BERRY DRUPE
An fleshy, indehiscent fruit that contains few to An fleshy, indehiscent fruit whose PERICARP is
many seeds (rarely one seed) and has a PERICARP differentiated into exo-, meso-, and endo-carp
that is entirely fleshy.
DRUPELET
A small drupe (e.g. segments of a raspberry)

BERRY DRUPE DRUPELET

MORPHOLOGY-BASED: DRY AND DEHISCENT FRUITS

CAPSULE LOCULICIDAL CAPSULE


A dry, dehiscent fruit that is derived from a Dehisces directly into the locule
syncarpous gynoecium and contains few to many
seeds. Capsules are the most common and PORICIDAL CAPSULE
variable of all fruit types. Dehisces by apical pores (e.g. Papaveraceae)

CIRCUMSCISSLE CAPSULE (PYXIS) SEPTICIDAL CAPSULE


Top comes off (like a lid) to release seeds (e.g. Dehisces along the septa
Portulaceae)
FOLLICLE
DENTICIAL CAPSULE A dry, dehiscent fruit that is derived from a
Dehisces by apical teeth (e.g. Caryophyllaceae) monocarpous or apocarpous gynoecium and
opens along a single suture

CIRCUMSCISSLE CAPSULE DENTICIDAL LOCULICIDAL SEPTICIDAL PORICIDAL FOLLICLE


(PYXIS) CAPSULE CAPSULE CAPSULE CAPSULE

Lab 4 - Fruit Terminology 123


MORPHOLOGY-BASED: DRY AND INDEHISCENT FRUITS

ACHENE SAMARA
A dry, indehiscent fruit that is derived from variable A winged ACHENE
gynoecium types and is small and single-seeded
(seed free from PERICARP) SAMAROID SCHIZOCARP
A winged SCHIZOCARP (e.g. maple)
NUT
A dry, indehiscent fruit that is large and single- SCHIZOCARP
seeded with a thick, bony PERICARP and often with A dry, indehiscent fruit that is derived from a
an INVOLUCRE (e.g. Fagaceae) syncarpous gynoecium that splits into two or more
one-seeded segments (MERICARPS)
NUTLET
A small NUT that does not possess an INVOLUCRE
(e.g. Lamiaceae)

ACHENE

NUT

NUTLET SAMARA SAMAROID SCHIZOCARP SCHIZOCARP

TAXONOMY-BASED FRUIT TYPES

CARYOPSIS LEGUME
POACEAE (GRASS FAMILY) FABACEAE (BEAN FAMILY)
A dry, dehiscent fruit with the seed coat fused to A dry, dehiscent fruit derived from a monocarpous
the pericarp (e.g. corn, wheat, rye, barley) gynoecium that usually opens along two sutures
(e.g. beans, peas)
CYPSELA (pl. CYPSELAE)
ASTERACEAE (ASTER FAMILY) PEPO
An achene fruit that usually has an adnate calyx CUCURBITACEAE (CUCUMBER FAMILY)
A fleshy, indehiscent, many-seeded fruit with a
HESPERDIUM thick rind that develops from an inferior ovary (e.g.
RUTACEAE (CITRUS FAMILY) pumpkin, squash, cucumber)
A fleshy, berry-like fruit with a leathery rind that
develops from a superior ovary (e.g. orange, lime)

CARYOPSIS CYPSELA HESPERIDIUM LEGUME PEPO

124 Integrative Biology 335: Plant Systematics


TAXONOMY-BASED FRUIT TYPES (CONT.)

POME SYCONIUM
ROSACEAE: MALOIDEAE (APPLE SUBFAMILY) MORACEAE (MULBERRY FAMILY)
A fleshy, indehiscent fruit derived from a (ONLY IN THE FICUS GENUS)
syncarpous gynoecium with an inferior ovary. A multiple fruit of many tiny drupes that are borne
Most of a pome’s flesh comes from the enlarged, inside of a hollow, inverted receptacle
fleshy hypanthium that grows up around the ovary
(i.e. apple, pear, quince)

SILIQUE AND SILICLE


BRASSICACEAE (MUSTARD FAMILY)
Dry, dehiscent fruits that have 2 valves which
separate from the persistent REPLUM (and
placentae). SILIQUES are typically more than twice
as long as wide, while SILICLES are typically less
than twice as long as wide

POME SILICLE SILIQUE SYCONIUM

SPECIAL FRUIT TYPES

ACCESSORY FRUIT
A fruit that has tissue other than, or in addition
to, the ovary that enlarges and may become
fleshy (e.g. strawberry, fig)

AGGREGATE FRUIT
A fruit that is the product of a single flower
with an apocarpous gynoecium (has multiple ACCESSORY FRUIT
distinct carpels). The ovary of each carpel
matures into a fruit (e.g. magnolia, tuliptree,
pawpaw, blackberry)

MULTIPLE FRUIT
A fruit that is the product of several separate
flowers in an inflorescence. (e.g. pineapple, MULTIPLE FRUIT
mulberry, sycamore, sweetgum) Other floral
structures (i.e., accessory tissue) may become
AGGREGATE FRUIT (e.g. OF DRUPELETS)
fleshy as well (e.g. strawberry)

Lab 4 - Fruit Terminology 125


MISCELLANEOUS FRUIT TERMINOLOGY

ARIL
A fleshy, often brightly colored seed cover
which arises from the funiculus and can
either partially or entirely cover a seed (e.g.
Magnoliaceae, Papaveraceae)

CARPOPHORE
A slender prolongation of the receptacle
forming a central axis between two carpels
(e.g. Apiaceae)

COMA
A tuft of hairs, especially on the tip of a seed
(e.g. Asclepiadaceae)

ELAIOSOME
Fleshy structures attached to seeds that are
rich in lipids and proteins (e.g. Papaveraceae)

ENDOCARP
The inner layer of a fruit’s PERICARP

EXOCARP
The outer layer of a fruit’s PERICARP

INVOLUCRE
A whorl of bracts subtending a flower cluster
(e.g. Fagaceae)

MESOCARP
The middle layer of a fruit’s PERICARP

PERICARP
The wall of a fruit

PLEUROGRAM
A groove on the surface of a seed (e.g.
Fabaceae)

REPLUM
Thin, papery partition between the two valves
of SILIQUES and SILICLES (e.g. Brassicaceae)

126 Integrative Biology 335: Plant Systematics


LABORATORY 5
MAGNOLIACEAE, RANUNCULACEAE AND
PAPAVERACEAE

PHYLOGENETIC RELATIONSHIPS

Enormous progress has been made recently in understanding phylogenetic relationships among flowering plants
(angiosperms). Such progress has proceeded so rapidly that many taxonomy texts are out of date very soon after
they are published! These new findings are having a major impact on our interpretation of angiosperm evoution and
the factors that account for the tremendous success of these plants.

Until rather recently, most students of angiosperm evolution considered those plants in the subclass “Magnoliidae”
as being among the first flowering plants to evolve. The “Magnoliidae” display an impressive range of morphological
variation, from large woody plants to small herbaceous ones. Some have large flowers with many spirally arranged
parts, while others have tiny flowers with few parts. Recent studies, however, have shown that the subclass
“Magnoliidae” is highly paraphyletic.

Recent studies have concluded that the first flowering plants to split from modern angiosperms were Amborella
trichopoda (a shrubby plant from the island of New Caledonia with small flowers and few spirally arranged parts)
and those belonging to the order Nymphaeales (water lilies). The Austrobaileyales form another very early branch
of angiosperm evolution. These results were surprising and led to the abandonment of subclass “Magnoliidae” as
the most basally branching lineage of flowering plants. Amborellaceae, Nymphaeaceae (the only family in the order
Nymphaeales) and Austrobaileyales are considered “basal families”. These three lineages subtend a clade including
all the rest of the flowering plants, which are referred to as the core angiosperms. We’ll have more to say about these
basal relationships, and the evolutionary implications of these results, in our lecture on the “Origin of Angiosperms”.

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 127


Within the core angiosperm clade, relationships
are still poorly resolved. The major clades that
are well-supported (by numerous independent
studies) include: (1) the Magnoliid complex or
clade, which includes the family Magnoliaceae,
(2) the Monocots, including such plants as
grasses, orchids, palms, lilies and bananas,
and (3) the Eudicots (or Tricolpates). The
Eudicots contain all other “dicots” not
contained within the Magnoliid clade. These
plants were originally called the tricolpate
clade, in reference to the main morphological
character marking this group - namely, the
occurrence of pollen grains with three colpi,
or germinal furrows. The Eudicots comprise
the early diverging “Basal Tricolpate” lineages,
containing such families as Ranunculaceae
and Papaveraceae, and the Core Tricolpate
clade.

This view of angiosperm relationships is quite


different from standard classifications in which
the flowering plants are divided into two major
groups, the monocots and “dicots”, and where
the subclass “Magnoliidae” is the earliest
diverging group of flowering plants. Both
“dicots” and “Magnoliidae” are paraphyletic,
thus these artificial groups are abandoned in
modern classifications. In their places, major
new clades have been identified, such as the
Eudicots and the Magnoliid complex.

The Magnoliid complex is made up of


Orders Magnoliales, Laurales, Canellales
and Piperales. The family Magnoliaceae
is monophyletic and is one of six families
comprising the Order Magnoliales.

To the left is a phylogeny of the Eudicots (or


Tricolpate clade of Angiosperms). The family
Ranunculaceae (Order Ranunculales) is a
member of the Basal Tricolpate group. Also
included in the Order Ranunculales are the families Papaveraceae and, in earlier treatments, the Fumariaceae. In
modern systems of classification, the Fumariaceae are submerged within the Papaveraceae.

128 Integrative Biology 335: Plant Systematics


MAGNOLIACEAE
magnolia family ♂ Ca3(x) Co6(x) A∞ G∞

Magnoliid Complex (Magnoliid Clade)

FLORAL CHARACTERISTICS

▪▪ Apocarpous gynoecium
▪▪ Many floral series are spirally arranged
▪▪ No connation or adnation of floral series
▪▪ LAMINAR STAMENS

Magnolia × soulangeana Soul. (Saucer Magnolia)

The species Magnolia × soulangeana Soul. is named


after Chevalier Soulange-Bodin who originally created the
hybrid (cross between Magnolia denudata and M. liliflora)
in his garden. Notice the use of the “×” in the scientific
name to indicate the tree’s hybrid origin. Its inflorescence
consists of solitary (terminal) flowers (Figure 5.1) that
are purplish or rose-colored outside, white within. While
observing the floral parts as instructed below, make sure
to check for connation and adnation.

1. Observe a flower. What is its symmetry?

2. CALYX: Look for the sepals at the base of the flower


(Figure 5.2). (Hint: Sepals are smaller than petals.)
Figure 5.1 Flower bud of Magnolia x soulangeana. How many are there?

3. COROLLA: Two series (or whorls) of petals are located


above the sepals. The first whorl of petals alternates
with the sepals, while the second whorl of petals
alternates with the petals below and are arranged
opposite the sepals. How many petals make up the
corolla?

Figure 5.2 Side view of a Magnolia x soulangeana


‘Andre LeRoy’ flower. The arrow points to
one of the 3 petal-like sepals.

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 129


4. ANDROECIUM: Observe the spirally arranged LAMINAR
STAMENS on the elongated receptacle. How many
stamens are there (many or few) and why is their
morphology considered to be ancestral?

5. GYNOECIUM: Check out the numerous spirally


arranged carpels (Figure 5.3). Although the carpels
may appear united when young, they are perceptibly
separate (distinct) at maturity. (To see this, observe
the dried fruit.) What is the term used to describe this
type of gynoecium? Is the ovary position superior or
inferior and what is the insertion type?

6. PORTFOLIO DRAWING: Draw a floral diagram of a


longitudinal section (L.S) of Magnolia × soulangeana,
Figure 5.3 Magnolia x soulangeana flower with several making sure to include its floral formula. Follow the
sepals, petals, and stamens removed. Note
format as instructed in Lab 3: Floral Terminology.
the spirally arranged stamens with laminar anthers and the
numerous distinct carpels on an elongated receptacle.

Liriodendron tulipifera L. (Tuliptree)

Examine a L. tulipifera flower if they are available. While


the color is very different from that of the Magnolia flower
you just dissected, note that the structure is essentially
the same, with many spirally arranged stamens, and a
gynoecium composed of many distinct carpels on an
elongate receptacle (Figure 5.4). The brown “hairs” on the
gynoecium are the styles/stigmas.

Figure 5.4 Longitudinal section of a Liriodendron


tulipifera, or tulip tree, flower.

130 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Aggregate fruits of follicles or samaras

Magnolia grandiflora L. (Southern Magnolia)


Liriodendron tulipifera L. (Tuliptree)

Observe the aggregate fruit of follicles produced by M.


grandiflora. This fruit is formed through elongation of the
receptacle and the development of the each of the carpels
into a follicle (Figure 5.5). Additionally, its seeds are
covered with red ARILS and remain attached to the follicle
by a thread-like FUNICULUS for some time after dehiscence.
Alternatively, L. tulipifera produces an aggregate fruit of
samaras (Figure 5.6). Based on their fruit morphology,
what do you suppose is the mechanism utilized by each
species for seed dispersal?
Figure 5.5 Magnolia grandiflora flower and fruit.

How can you tell that the fruits on display were derived
from a single flower?

Figure 5.6 Liriodendron tulipifera flower and fruit.

Box 1 HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Has stipules
▪▪ Stipular scars surrounding twigs

Michelia spp. (Michelia)

Michelia is related to our much larger magnolias (Magnolia


spp.) and tuliptrees (Liriodendron tulipifera). To review
your vegetative terminology, observe a Michelia leaf,
sketch it in Box 1 and then add labels describing the leaf
shape, apex, base, venation and margin. Don’t forget to
include the axillary bud!

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 131


Magnolia acuminata (L.) L. (Cucumber-tree)
Liriodendron tulipifera L. (Tuliptree)

Using the “Dichotomous Key for Woody Plants” in Lab 1:


Woody Twig Terminology, determine which twig is from
a Magnolia and which is from a Liriodendron tree. Which
characteristics set them apart?

One of the key features used to separate these


Magnoliaceae species from other woody species in the
dichotomous key is the occurrence of STIPULES covering
the flower bud. In the spring the stipules dehisce leaving
a STIPULE SCAR that surrounds the twig. What purpose do
you think stipules serve?

Locate the STIPULES, STIPULE SCARS (on older branches),


LEAF SCARS and BUNDLE SCARS on the branches of the live
specimens provided and then label those same structures
in Figure 5.7.

Figure 5.7 Winter twig of a Magnoliaceae species.


Notice the distinctive stipules and how the
stipular scars completely encircle the twig.

132 Integrative Biology 335: Plant Systematics



RANUNCULACEAE
buttercup family ♂ Ca4-∞ Co0-∞ A∞ G∞

Eudicots: Basal Eudicots

FLORAL CHARACTERISTICS

▪▪ Parts spirally arranged


▪▪ Apocarpous gynoecium
▪▪ Many marginal ovules or 1 basal ovule
▪▪ Stamens NOT laminar

Delphinium spp. (Larkspur)

To observe the floral morphology characteristic of the


Ranunculaceae family you will examine and dissect a
Delphinium flower. There are over 250 species in this
genus (commonly called larkspurs), including many that
are showy and popular garden plants. These plants
Figure 5.8 A Delphinium flower. Notice its 5 sepals, one contain alkaloids and are usually quite toxic.
of which is a nectar spur.

1. Determine the Delphinium plant’s inflorescence type


as well as your flower’s symmetry.

2. CALYX: Note that the posterior sepal is prolonged


into a long NECTAR SPUR and two other sepals form
a landing platform for insects (Figures 5.8 and 5.9).
Also, observe the prominent color of these showy
sepals. How many sepals are there?

3. COROLLA: The upper pair of petals also have spurs


that project backward and down into the sepal spur
where there is nectar. The remaining two petals, if
present, are often crowded in the throat and are small
and short-clawed (these are sometimes called the
“bee”). How many petals does your flower have?

4. ANDROECIUM: Typically the stamens are spirally


arranged in Ranunculaceae flowers, but it may be
Figure 5.9 A flower of Delphinium in longitudinal section.
hard to detect in this species. How many stamens are
there?

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 133


5. GYNOECIUM: What type of gynoecium does the
Box 2
Delphinium flower possess and how many carpels
make it up? Sketch out the placentation type of the
flower in Box 2.

6. Write out the floral formula for the Delphinium flower


you dissected in the space below.

Aconitum spp. (Monkshood)

Aconitum flowers contain powerful poisons, and drugs


are derived from the roots and leaves of certain species
(Figure 5.10). Observe its flowers and compare them
to the following description: Flowers hypogynous,
zygomorphic (most members of Ranunculaceae have
actinomorphic symmetry), and all parts are free and
distinct. There are 5 sepals; the upper sepal (called the
helmet) is the largest and is strongly arched or hooded,
its tip prolonged forward and downward into a short beak.
The upper two petals are concealed under the helmet,
clawed and nectariferous at the tip. The lower three petals
are absent or vestigial. Stamens are numerous, while
Figure 5.10 An Aconitum flower.
the apocarpous gynoecium is made up of 3 to 5 distinct
carpels. Aconitum produces an aggregate fruit of follicles.

Helleborus spp. (Hellebore)

Check out the dissected flower of this species as well as


the provided photos (Figure 5.11). Species in this genus
have solitary, actinomorphic flowers subtended by leafy
bracts, with five green or petaloid sepals and petals
modified into small, tube-like nectaries. What do you
suppose is the function of the nectaries?

Figure 5.11 A Helleborus flower with its small, tube-like


nectaries (modified petals).

134 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruits are follicles or achenes

Aquilegia spp. (Columbines)

The genus name Aquilegia comes from the Latin word


for eagle, “aquila,” since its characteristic spurred petals
resemble an eagle’s claw. Additionally, the English
common name, columbine, alludes to the Latin word for
dove, “columba,” because the spurred petals also appear
to some like a group of doves drinking (Figure 5.12). What
is Aquilegia’s fruit type (be specific)?
Figure 5.12 Flower longitudinal section (left) and
developing follicles (right) of Aquilegia
canadensis. Two of the carpels (developing follicles) have
been split open to show their marginal placentation.

Clematis spp. (Clematis)

The flowers of this plant have no petals (the showy


perianth parts are sepals), numerous stamens and
carpels. Based on the provided herbarium specimens and
photographs (Figure 5.13), what fruit type do Clematis
species produce and what gynoecium type do these fruit
arise from? Also, what are the vegetative structures that
allow this plant to climb?

Figure 5.13 Fruit of Clematis virginiana, commonly called


Virgin’s Bower.

Actaea pachypoda Elliot (Doll’s Eyes)

Observe the photographs and herbarium specimen of this


species (Figure 5.14). Actaea pachypoda produces a fruit
type that is rather rare in the Ranunculaceae family. What
is this fruit type called? What seed dispersal mechanism
do you think this species uses?

Figure 5.14 Developing infructescence (top) and fruit


cross section (bottom) of Actaea pachypoda.

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 135


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Perennial or annual herbs


▪▪ Without stipules

Anemone patens L. (American Pasqueflower)


Hepatica nobilis Shreb. (Liverleaf)

Like A. patens and H. nobilis (a common woodland


wildflower in Illinois), the habit of most Ranunculaceae
species is an herb (Figures 5.15 and 5.16). (Note:
remember that Magnoliaceae are ALL woody). These
herbs can be either annual or perennial. What does annual
and perennial mean?

Figure 5.15 Habit of Anemone patens, commonly called


American Pasqueflower.

Figure 5.16 Habit of Hepatica nobilis, commonly called


Liverleaf.

136 Integrative Biology 335: Plant Systematics


PAPAVERACEAE
poppy family ♂ Ca2(4) Co4-8 A6 or ∞ G 2-∞

Eudicots: Basal Eudicots

FLORAL CHARACTERISTICS

▪▪ Two sepals are CAUDUCOUS


▪▪ Petals sometimes crumpled in bud
▪▪ Syncarpous gynoecium
▪▪ Many parietal ovules

Argemone mexicana L. (Mexican Prickly Poppy)

To observe the floral morphology characteristic of the


Papaveraceae family, you will examine and dissect an
A. mexicana flower (Figure 5.17). The genus name,
Argemone, comes from the Greek “argema,” or cataract,
because the plant was originally thought to cure eye
cataracts.

1. What is the floral symmetry of the flower?


Figure 5.17 The prickly poppy, Argemone mexicana, is
appropriately named as the whole plant is
covered with prickles, even the sepals before they fall off
(white arrow pointing to prickly sepals.)

2. CALYX: Can you find the sepals on your specimen?


Don’t be surprised if you can’t. In this family the
sepals are CAUDUCOUS (that is they fall off as the
flower opens). To locate the sepals, you need to
look at an unopened flower bud (the sepals will fully
enclose the bud.)

3. COROLLA: Papaveraceae flowers generally have


twice as many petals as sepals. How many petals
does your flower have?

4. ANDROECIUM: Note the arrangement of the numerous


stamens (Figure 5.18). Like Magnoliaceae and
Ranunculaceae, the stamens are spirally arranged.

Figure 5.18 A flower of Stylophorum diphyllum in


longitudinal section.

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 137


5. GYNOECIUM: Is the ovary position superior or inferior?
Box 3
Also, what type of gynoecium does the flower possess
and how many carpels make it up? (HINT: Count the
stigma lobes.) Now draw a cross section (X.S.) of the
ovary in Box 3 to show its placentation type.

6. PORTFOLIO DRAWING: Draw a floral diagram of a


longitudinal section of Argemone as well as a cross
section (X.S.) of its ovary, making sure to include its
floral formula. Follow the format as instructed in Lab
3: Floral Terminology.

Dicentra spp. (Bleeding Heart)

The genus Dicentra belongs to Fumerioideae, a subfamily


of Papaveraceae. Based on the live Dicentra specimen
and provided photographs (Figure 5.19), what are two
floral characteristics that set this subfamily apart from
other Papaveraceae species?

Figure 5.19 Flowers of Dicentra canadensis. These


flowers are zygomorphic and possess two
very different shaped petals - (2) large ones that have nectar
spurs at the base and (2) narrow ones that have expanded
tips to enclose the anthers and stigma.

FRUIT TYPE

▪▪ Capsules (Poricidal in Papaver)


▪▪ Seeds have ELAIOSOMES for ant dispersal

Papaver spp. (Poppies)


Glaucium spp. (Horned Poppy)

The fruits of Papaver and Glaucium display a “shaker”


method of dispersing seed (Figure 5.20). They are on
long stems that wave in the wind, shaking the seeds out
through the pores. What specific type of fruit do these
species produce? How does it dehisce?

Figure 5.20 Poricidal capsule fruit of Papaver species. It


dehisces by small pores around the top.

138 Integrative Biology 335: Plant Systematics


Stylophorum diphyllum (Michx.) Nutt.
(Celandine poppy)

Take a close look at the seeds produced by S. diphyllum


with their distinctive ELAIOSOMES (fleshy structures
attached to seeds that are rich in lipids and proteins;
Figure 5.21). Due to the elaiosomes’ nutritional value, ants
will come gather the seeds and carry them far from the
parent plant to their nest as food for ant larvae. After the
larvae have consumed the elaiosome, the undamaged
seed is moved to the ants’ disposal area. Typically the
soil found in disposal areas is rich in nutrients and thus
good for seed germination. This is a great example of a
mutualistic relationship!

Figure 5.21 Close-up of seeds of Stylophorum diphyllum.


Note the pale colored elaiosome on each
seed that attracts ants to carry the seeds away from the
parent plant.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Perennial herbs
▪▪ Sap milky or colored

Check out the wide range of milky or colored sap that


herbaceous species in the Papaveraceae family exhibit.
Some species’ sap contains alkaloids.

Chelodonium majus L. (Celandine)

C. majus produces an umbel-like inflorescence of yellow


flowers and will exude yellow sap when its stem is cut.

Figure 5.22 Herbaceous habit of Sanguinaria canadensis,


a woodland wildflower. Bocconia frutescens L. (Tree Poppy or Parrotweed)

This member of the Papaveraceae family is an oddity


in that it is a tree whereas most other species are
herbaceous. B. frutescens does, however, produce orange
sap that can be seen when some of the bark on a branch
is scraped away.

Sanguinaria canadensis L. (Bloodroot)

“Bloodroot”, the common name of S. canadensis, is


very fitting due to the red sap that is stored in the plant’s
RHIZOME (Figures 5.22 and 5.23). The red sap is also
present in the stem and thus is a good feature to use for
identification in the field.

Dicentra spectabilis (L.) Lem. (Bleeding-heart)

A common garden plant, D. spectabilis, or bleeding


heart, produces clear sap like most members of the
Figure 5.23 Cut stems of Sanguinaria canadensis oozing Fumarioideae subfamily.
orange-red sap.

Lab 5 - Magnoliaceae, Ranunculaceae and Papaveraceae 139


140 Integrative Biology 335: Plant Systematics
LABORATORY 6
HAMAMELIDACEAE, MORACEAE, FAGACEAE AND
BETULACEAE

PHYLOGENETIC RELATIONSHIPS

Hamamelidaceae is one of thirteen families


belonging to the order Saxifragales. The
family Hamamelidaceae is highly variable and
its monophyly has been questioned. Some
systematists now separate one common tree
species, Liquidambar styraciflua L. (Sweet Gum),
into its own family, Altingiaceae.

Moraceae is treated as its own distinct family


by Judd et al., but as a subfamily of Urticaceae
by Zomlefer. It is classified in the order Rosales
of the Rosid clade. The Rosids are a rather
heterogeneous grouping of orders that are
supported solely by molecular evidence. The
monophyly of Moraceae is supported only
by molecular evidence. These plants are
characterized by milky sap. Major genera include
Ficus (figs), Morus (mulberries), Maclura (osage
orange), Artocarpus (jackfruit, breadfruit) and
Dorstenia.

Fagaceae and Betulaceae are members of the


order Fagales in the Rosid clade. The order Fagales consists of 8 families and about 1,115 species. Traditionally,
the Fagaceae and Betulaceae families were united with the family Hamamelidaceae in the subclass “Hamamelidae”.
Recent evidence suggests the Hamamelidaceae and Fagales are not at all closely related. The “Hamamelidae”, as
traditionally defined to include the four families covered in today’s lab, are highly polyphyletic.

Lab 6 - Hamamelidaceae, Moraceae, Fagaceae and Betulaceae 141


HAMAMELIDACEAE ♂ Ca 4-5 Co4-5(10) A4-5 OR

witch hazel family
G②
Eudicots: Core Eudicot ♂ Ca 4-5 Co4-5(10) A4-5 G②

FLORAL CHARACTERISTICS

▪▪ 4-merous flowers with strap-like petals


▪▪ STAMINODES present in Hamamelis
▪▪ Anthers open by flaps in Hamamelis

Hamamelis mollis Oliv. (Witch Hazel)

To observe the floral morphology characteristic of


Hamamelidaceae, you will examine a H. mollis flower.

1. Examine a flower under a dissecting scope. What is


its symmetry?

2. CALYX: Notice at the base of the flower that there are


several bracts. Inside the layer of bracts you should
find the sepals. How many sepals are in the calyx?
Figure 6.1 Cluster of Hamamelis flowers.

3. COROLLA: The corolla is made up of 4 linear, strap-


shaped petals (Figures 6.1 and 6.2).

4. ANDROECIUM: Four stamens make up the androecium


of this flower. Their anthers open by flaps to release
pollen. Also note the STAMINODES located at the base
of the petals. What are their purpose?

5. GYNOECIUM: The ovary is half-inferior and the


gynoecium is surrounded by a dense pubescence.
Make a longitudinal section of this flower between the
two styles (the 2 carpels are basally connate). If done
successfully, you should see a solitary, pendulous
ovule in each of the 2 locules.

6. PORTFOLIO DRAWING: Draw a floral diagram of a


Figure 6.2 Longitudinal section of a Hamamelis flower. longitudinal section of H. mollis, making sure to
include the floral formula and label all the floral
parts. Follow the format as instructed in Lab 3: Floral
Terminology.

142 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruit is a woody loculicidal capsule

Hamamelis mollis Oliv. (Witch Hazel)

The ovary of H. mollis, like other Hamamelidaceae


species, matures into a loculicidal capsule bearing two
black seeds (these seeds are forcibly discharged when
the fruit is ripe; Figure 6.3). How can you tell the capsule is
loculicidal and not septicidal?

Liquidambar styraciflua L. (Sweet Gum)

Observe the L. styraciflua fruits on display (Figure 6.4).


Figure 6.3 Mature capsule of Hamamelis vernalis. Formerly in Hamamelidaceae, this species is now placed
in its own family, Altingiaceae. The flowers of Altingiaceae
are wind pollinated and the seeds (which are often
winged) are wind-dispersed. What type of fruit is this (be
specific)?

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Toothed or lobed leaves


▪▪ Stipules present
▪▪ Stellate hairs often present
▪▪ Often colorful in autumn

Figure 6.4 Multiple fruit of loculicidal capsules produced


by Liquidambar styraciflua.
Liquidambar styraciflua L. (Sweet Gum)

The leaves of L. styraciflua are very distinctive, however


in the winter one must rely on twig morphology. Using the
“Dichotomous Key for Woody Plants” in Lab 1, key out the
winter twig of L. styraciflua.

Hamamelis vernalis Sarg. (Ozark Witchhazel)

Like many members of the Hamamelidaceae family, H.


vernalis, also called spring or vernal witchhazel, has
alternate, simple, rather broad leaves with coarsely
toothed margins (Figure 6.5). If this photograph were in
color, you could see the vibrant yellow color the leaves
Figure 6.5 Fruits and leaves of Hamamelis vernalis. turn in the autumn.

Lab 6 - Hamamelidaceae, Moraceae, Fagaceae and Betulaceae 143


MORACEAE
mulberry family
♂ Ca4 Co0 A4 G0

Eudicots: Rosid Clade ♀ Ca4 Co0 A0 G②

FLORAL CHARACTERISTICS

▪▪ SYCONIUM inflorescence
▪▪ Imperfect flowers

Ficus diversifolia Blume. (Mistletoe Fig)

The fig is a native of Asia Minor that was imported into


the Mediterranean area and used by the Egyptians 6000
years ago. Figs were first introduced into North America
around 1600 but commercial cultivation was not begun in
this country until 1900. F. diversifolia produces a SYCONIUM
inflorescence (a SYCONIUM is also a fruit type). Its small
unisexual and apetalous flowers are located inside the
SYCONIUM (Figure 6.6).

Longitudinal section of a Ficus diversifolia


Carpellate Flowers
Figure 6.6
SYCONIUM.
Note the fertile, white carpellate flowers at the base of the
inflorescence with their long styles and stigmas and their
orange, fleshy sepals. The ovary is usually unilocular with
a single, apical ovule.

Staminate Flowers

Covering the pore at the top of the synconium are


numerous scales and below these are the staminate
flowers. Typically staminate flowers will have few and
often only one stamen per flower. Depending upon your
synconium, male flowers may not be obvious.

PORTFOLIO DRAWING: Draw a floral diagram of a


longitudinal section of your F. diversifolia SYCONIUM,
making sure to include the floral formula and label all the
floral parts. Follow the format as instructed in Lab 3: Floral
Terminology.

Dorstenia carnulosa Dewild. (Dorstenia)

Compare the inflorescence of Dorstenia (Figure 6.7) to


that of Ficus. How are they different?
Figure 6.7 Dorstenia carnulosa inflorescence.

144 Integrative Biology 335: Plant Systematics


Box 1 FRUIT TYPE

▪▪ Achenes or small drupes


▪▪ Sometimes multiple fruits with accessory tissue
(SYCONIUM)

Ficus carica L. (Smyrna Fig)

The fig (SYCONIUM) is a peculiar fruit, with its fleshy


receptacle surrounding the actual fruits (small drupes).
Thus, a SYCONIUM could be considered a multiple
fruit with accessory tissue. The common fig develops
parthenocarpically (without fertilization of the flowers),
and the fruits contain no embryos. However, the Smyrna
fig must be pollinated by wasps, Ficus species’ typical
pollinator, in order to develop fruit. This particular variety
produces only female flowers and depends on the wild
caprifig tree for pollination. The grower of Smyrna figs
must tie caprifigs containing wasp eggs to the branches
of his Smyrna trees. The eggs hatch, the wasps develop,
they exit the caprifig and pick up some pollen and then
visit Smyrna figs, pollinating them in the process. Dissect
a Smyrna fig and try to find a wasp inside. Yummy! Now,
create a simple diagram in Box 1 of the wasp-fig life cycle
based on your lecture material.

Morus spp. (Mulberries)

Morus species have catkin inflorescences that, when


pollinated, develop into multiple fruits of drupelets (Figure
6.8). You might notice that mulberries look very similar
to raspberries (Rubus idaeus (Rosaceae)) superficially;
however raspberries are classified as aggregate fruits of
drupes. What is the difference between these two fruit
types? (HINT: Think flower number and gynoecium type.)

Figure 6.8 Morus rubra’s multiple fruit of drupelets.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Trees and shrubs


▪▪ Milky sap

Maclura pomifera (Raf.) C.K. Schneid. (Osage Orange)

The habit of M. pomifera is a tree, and it can be easily


Figure 6.9 Maclura pomifera fruit.
recognized by its orangish-brown bark, branches with
axillary spines, and multiple fruit of drupes that some

Lab 6 - Hamamelidaceae, Moraceae, Fagaceae and Betulaceae 145


people think resemble green brains (Figures 6.9 and
6.10). Key out the M. pomifera winter twig using the
“Dichotomous Key for Woody Plants” in Lab 1.

M. pomifera has separate male and female inflorescences


that are located on different plants. Based on this
information, what is the plant condition?

Figure 6.10 Fruit and trunk of a Maclura pomifera tree.

146 Integrative Biology 335: Plant Systematics


FAGACEAE ♂ Ca 4-7 Co0 A4-∞ G0
beech or oak family
♀ Ca 4-7 Co0 A0
Eudicots: Rosid Clade
G③

INFLORESCENCE TYPE

▪▪ Staminate flowers in catkins and carpellate flowers in few-


flowered clusters enclosed by an INVOLUCRE

Quercus spp. (Oaks)

Observe the Quercus woody twigs and try to identify


the STAMINATE CATKINS as well as the OLD and YOUNG
CARPELLATE FLOWERS (Figure 6.11). Label these
structures in the photograph below.

1. STAMINATE CATKINS: long and pendulous

2. OLD CARPELLATE FLOWERS: these flowers from the


previous year are now developing into ACORNS

3. YOUNG CARPELLATE FLOWERS: arranged in few


Figure 6.11 Quercus macrocarpa inflorescences. flowered clusters enclosed by an INVOLUCRE

Lab 6 - Hamamelidaceae, Moraceae, Fagaceae and Betulaceae 147


FLORAL CHARACTERISTICS

▪▪ Gynoecium of 3 connate carpels with an inferior ovary

Quercus spp. (Oaks)

To observe the floral morphology characteristic of


Fagaceae, you will examine staminate and carpellate
Quercus flowers. Quercus and all Fagaceae flowers are
imperfect and located on the same plant. Therefore, what
is the plant condition?

Figure 6.12 Female flowers of Quercus rubra.


Carpellate Flowers

Examine the few flowered clusters of the new (this year’s)


carpellate flowers and the INVOLUCRE that surrounds them
(Figure 6.12). You’ll notice that the carpellate flowers
have an inconspicuous, partially connate calyx, but lack
a corolla and an androecium. Identify the stigma, style
and ovary of the syncarpous gynoecium. Based on the
number of stigmas, how many connate carpels form this
gynoecium? Is the ovary position superior or inferior? Also,
what is the insertion type?

Staminate Flowers

As previously mentioned, the small staminate flowers are


arranged into the long, dangling CATKINS characteristic of
the Fagaceae family. Staminate flowers possess a scale-
Figure 6.13 1-year old female Quercus rubra flowers
like calyx of 4 to 7 connate sepals, but lack a corolla and a
developing into acorns. gynoecium (Figure 6.14). Their androecium is made up of
4 to many un-fused stamens. What is the floral part fusion
term that could be used to describe these stamens?

Figure 6.14 Catkin of male Quercus pumila flowers.

148 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruit is a nut surrounded by an INVOLUCRE

Castanea spp. (Chestnuts)


Fagus sylvatum L. (European Beech)
Fagus grandifolia Ehrh. (American Beech)
Quercus spp. (Oaks)

In Fagaceae, the ovary of each flower within an


INVOLUCRE develops into a nut. Thus, in Castanea,
Figure 6.15 Fagus grandifolia (left) and Fagus sylvatica
(right) nuts with surrounding involucres. Fagus and Quercus species there are 3, 2 and 1 nut(s),
respectively, enclosed in each INVOLUCRE (Figures 6.15
and 6.16). Observe the diversity in INVOLUCRE morphology
among the 3 genera.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Trees or sometimes shrubs

Quercus macrocarpa Michx. (Bur Oak)


Figure 6.16 Acorns of Quercus palustris.
Quercus rubra L. (Northern Red Oak)
Fagus sylvatica L. (European beech)

Identify the species listed above using the “Dichotomous


Key for Woody Plants” in Lab 1. Then label the key twig
structures in Figure 6.17.

Figure 6.17 Quercus palustris winter twigs.

Lab 6 - Hamamelidaceae, Moraceae, Fagaceae and Betulaceae 149


BETULACEAE ♂ Ca 0-6 Co0 A2-18 G0
birch family
♀ Ca 0-6 Co0 A0
Eudicots: Rosid Clade
G②

INFLORESCENCE TYPE

▪▪ Staminate and carpellate flowers clustered into separate


CATKINS

Alnus glutinosa (L.) Gaertn. (European Alder)

Observe the woody twigs that have been collected


for A. glutinosa and try to identify the three CATKIN
inflorescences types listed below as well as the twig’s
VEGETATIVE BUDS. Then label those structures on the
photograph of A. glutinosa below.

1. STAMINATE CATKINS: these are long and pendulous


and may be releasing yellow pollen

2. OLD CARPELLATE CATKINS: these are found at the end


of the twig, are woody and resemble small pine cones

3. YOUNG CARPELLATE CATKINS: these are very small


and are found next to the long pendulous staminate
catkins
Figure 6.18 Catkins of American Hazelnut, also known
as Corylus americana. 4. VEGETATIVE BUDS

Figure 6.19 Betula pendula branch.

150 Integrative Biology 335: Plant Systematics


FLORAL CHARACTERISTICS

▪▪ Gynoecium of 2 connate carpels with an inferior ovary

Alnus glutinosa (L.) Gaertn. (Common or Black Alder)

To observe the floral morphology characteristic of


Betulaceae, you will examine staminate and carpellate A.
glutinosa flowers.

Carpellate Flowers

The carpellate catkins are very short, with each bract


Figure 6.20 Ostrya virginiana male and female catkins.
subtending two flowers (a CYMULE) that lack a calyx and
corolla (Figure 6.21). The flowers’ ovaries are inferior, bi-
carpellate and have two styles. Based on this, what type of
gynoecium do you think the flowers have?

Write out the formula for a female A. glutinosa flower given


the provided information.

Staminate Flowers

The staminate catkins are pendulous. Each bract subtends


3 tiny flowers (a CYMULE) and each flower consists of a
Figure 6.21 Female catkin of Alnus cordata. small 4-parted calyx and 4 stamens with short filaments.
There are no petals (corolla). The anthers are deeply
divided so that the pollen sacs are distinct. Observe a
portion of a staminate catkin under your dissecting scope
and identify the bracts and individual flowers with their
calyx and stamens. Write out the formula for a male A.
glutinosa flower given the provided information.

Figure 6.22 Male flowers of Betula pendula.

FRUIT TYPE

▪▪ Fruits are samaras and one-seeded nuts

Alnus glutinosa (L.) Gaertner (Black Alder)

Figure 6.23 Mature female catkin of Alnus glutinosa. The carpellate catkins of Alnus species are persistent, and

Lab 6 - Hamamelidaceae, Moraceae, Fagaceae and Betulaceae 151


become woody as they mature (Figure 6.23). The small
samara fruits contained within are gradually released with
movement of the mature, woody catkin.

Corylus americana Walter (American Hazelnut)

C. americana produces the other fruit type characteristic


of Betulaceae, a nut (Figure 6.24). Check out the provided
fruits, herbarium specimen and illustrations. What is
the name of the green structures enclosing these C.
americana nuts?

Figure 6.24 Fruits of Corylus americana.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit is a tree or a shrub


▪▪ Doubly serrate leaf margins

Betula pendula Roth (European White Birch)

Using the “Dichotomous Key for Woody Plants” in Lab 1,


be sure you can identify a winter twig of B. pendula.

Figure 6.25 Ostrya virginiana fruits and leaves.

152 Integrative Biology 335: Plant Systematics


LABORATORY 7
CACTACEAE, CARYOPHYLLACEAE, PORTULACEAE
AND POLYGONACEAE

PHYLOGENETIC RELATIONSHIPS

The Caryophyllid clade belongs within the


Core Eudicots (Core Tricolpates) clade. The
monophyly of the Caryophyllid clade is supported
by seed coat anatomy, anther wall development
and molecular data. However, their position,
whether within the Rosid complex or sister to the
Asterid clade is uncertain. The Caryophyllid clade
contains two large orders, Caryophyllales and
Polygonales.

The order Caryophyllales is clearly monophyletic,


as evidenced by their numerous, distinctive
synapomorphies, such as an embryo curved
around the seed, the presence of perisperm
instead of endosperm, free-central or basal
placentation, sometimes a single whorl of tepals,
a specialized type of plastid with a central protein
crystal, loss of the rpl2 intron (a major structural
rearrangement of the chloroplast genome),
and betalain pigments instead of anthocyanins
(except in the family Caryophyllaceae). Recent
cladistic analyses suggest that the anthocyanin-
possessing Caryophyllaceae evolved from
betalain-containing ancestors that lost their betalains and reacquired anthocyanins. These two changes are possibly
biochemically interrelated. Of course, the monophyly of the Caryophyllales is strongly supported by comparative
DNA sequence data. The Caryophyllales consists of 18 families and 8,600 species, with its major families including
Cactaceae, Portulacaceae and Caryophyllaceae.

The Caryophyllales are closely related to the order Polygonales; indeed, in many phylogenetic analysis, they are
sister taxa. The Angiosperm Phylogeny Group actually places Polygonales in the order Caryophyllales. Among the
10 families (about 2,050 species) included in Polygonales, are Polygonaceae and Plumbaginaceae as well as the
carnivorous plant families Droseraceae and Nepenthaceae. It is of interest to note that the pitcher-like leaves of the
Old World family Nepenthaceae are amazingly similar with those of the distantly related, New World carnivorous
family Sarraceniaceae. This an amazing case of convergent evolution!

Many of the families of the Caryophyllid clade were treated previously by Arthur Cronquist in the subclass
Caryophyllidae. Unlike most other subclasses of flowering plants, the Caryophyllidae are supported as monophyletic
through a variety of cladistic analyses of both morphological and molecular data.

Lab 7 - Cactaceae, Caryophyllaceae, Portulacaceae and Polygonaceae 153


CACTACEAE
cactus family ♂ Ca∞ Co∞ A∞

G 3-∞
Eudicots: Caryophyllid Clade

THREE SUBFAMILIES

▪▪ CACTOIDEAE: Stems generally not jointed and glochids


absent

▪▪ OPUNTIOIDEAE: Jointed stems and glochids (each stem-


joint represents a season’s growth)

▪▪ PERESKIOIDEAE: Leafy shrubs with scarcely succulent


stems

FLORAL CHARACTERISTICS

▪▪ Large and showy flowers


Flowers of Schlumbergera bridgesii. ▪▪ Ovary inferior and sunken into stem tissue
Figure 7.1

Schlumbergera spp. (CACTOIDEAE) (Schlumbergera)

To observe the floral morphology characteristic of


Cactaceae, you will examine and dissect a Schlumbergera
flower (either S. bridgesii (Lemaire) Loefgren., the
Christmas Cactus, or S. truncata (Haw.) Moran, the Crab
Cactus). This genus is native to Brazil and many cultivars
have been produced. In their native habitat, these plants
grow as epiphytes on trees. What is an epiphyte?

1. Observe a flower. What is its symmetry?

2. CALYX and COROLLA: The perianth elements increase


in size and gradually transition from sepals to petals
from the outermost to the innermost whorl (Figures
7.1 and 7.2). Since sepals and petals cannot be
distinguished, they are all called TEPALS. In this flower
a hypanthium is present (not shown in floral formula).
Look for it just above the ovary (it’s rather small).
What adnate parts form the hypanthium?

Figure 7.2 Longitudinal section of a Schlumbergera


bridgesii flower.

154 Integrative Biology 335: Plant Systematics


3. ANDROECIUM: Are there few or numerous stamens in
this flower?

4. GYNOECIUM: Note the inferior ovary sunken into the


stem tissue with its one locule. How many carpels are
fused to produce this syncarpous gynoecium? (HINT:
Count the number of stigma lobes.)

Figure 7.3 Flower of an Opuntia macrorhiza flower.

Opuntia macrorhiza Engelm. (OPUNTIOIDEAE)


(Twistspine Pricklypear)

O. macrorhiza is one of the three Opuntia species native


to Illinois (Figure 7.3). The other two are O. humifusa and
O. fragilis (an endangered species).

Observe Figure 7.4 depicting a longitudinal section of O.


macrorhiza. What is its insertion type?

FRUIT TYPE

▪▪ Fruit is a berry

Figure 7.4 Longitudinal section of an Opuntia


macrorhiza flower. Opuntia spp. (OPUNTIOIDEAE)

The fruits of Cactaceae are many-seeded berries (Figure


7.5). Since the ovary is often imbedded in stem tissue,
the berry is likewise. Hence, AREOLES with SPINES and/
or GLOCHIDS are often found on the outside of the berries.
The berries of some species of Opuntia are edible and
sold in supermarkets. However, be sure to remove the skin
and GLOCHIDS before trying to eat one!

Figure 7.5 Berry fruits of Opuntia.

Lab 7 - Cactaceae, Caryophyllaceae, Portulacaceae and Polygonaceae 155


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Stem succulents with BETALAIN pigments


▪▪ Leaves reduced or absent
▪▪ SPINES (modified leaves) produced from an AREOLE which
sometimes contains numerous GLOCHIDS

Echinocactus spp. (CACTOIDEAE)


Cephalocereus senilis (Haw.) Pfeiff. (CACTOIDEAE)

As you can see, Echinocactus and C. senilis (the old man


cactus) from Cactoideae are stem succulents and contain
numerous AREOLES and SPINES (Figure 7.6). Areoles
are small structures on the photosynthetic stems of cacti
that are usually interpreted as rudimentary shoots. Often
spines (reduced leaves borne on this shoot) are produced
at the areole.

Opuntia spp. (OPUNTIOIDEAE)

Species within the Opuntioideae subfamily also have


short yellowish or dark colored bristles called GLOCHIDS
in addition to their AREOLES and SPINES. GLOCHIDS (or
GLOCHIDIA) are actually a type of defense mechanism
since they are easily detached from the AREOLE when
brushed against by animals (such as humans) and
Figure 7.6 AREOLES and SPINES of Cactaceae. contain downward pointing barbs that force themselves
into the skin, causing great discomfort (definitely an
understatement!) (Figure 7.7). They are especially
abundant in the genus Opuntia. Draw and label AREOLES,
Box 1
SPINES and GLOCHIDS in Box 1 after observing the Opuntia
specimens on display.

Pereskia spp. (PERESKIOIDEAE)

Observe the vegetative characteristics of Pereskia. Why


are members of the subfamily Pereskioideae, such as
Pereskia, considered ancestral within Cactaceae?

Figure 7.7 A highly magnified photograph of a GLOCHID


taken with a scanning electron microscope.

156 Integrative Biology 335: Plant Systematics


CARYOPHYLLACEAE
pink family ♂ Ca 5 Co5 A10 G 2-5

Eudicots: Caryophyllid Clade

INFLORESCENCE TYPE

▪▪ CYME inflorescences

Saponaria officinalis L. (Bouncing Bet)

The basic inflorescence in the Caryophyllaceae is a CYME,


as seen here in S. officinalis (Figure 7.8). Remember that
a CYME is an example of a DETERMINATE INFLORESCENCE.
What does that mean?

Figure 7.8 Cyme inflorescence of Saponaria officinalis.

Box 2
Draw a cyme inflorescence in Box 2. Then label your
drawing with the following terms: terminal flower, lateral
flower, bracts, pedicel, peduncle.

FLORAL CHARACTERISTICS

▪▪ Petals often clawed at base and apically notched


▪▪ Ovary on a GYNOPHORE (i.e. ANDROGYNOPHORE), a stalk
that supports the Corolla, Androecium and Gynoecium

Dianthus spp. (Pinks)

To observe the floral morphology characteristic of


Caryophyllaceae, you will examine a Dianthus flower.

1. Examine a flower (Figure 7.9). What is its symmetry?

Figure 7.9 Flower of a Dianthus cultivar.

Lab 7 - Cactaceae, Caryophyllaceae, Portulacaceae and Polygonaceae 157


2. CALYX: The sepals in the calyx are fused to form a
ribbed tube. To determine the number of sepals, count
the number of calyx tube lobes.

3. COROLLA: The petals are clawed at their base, and


at the juncture of the CLAW and LIMB on the inner
surface are two APPENDAGES (Figure 7.10). How
many petals are there?

4. ANDROECIUM: How many stamens make up the


androecium? Are they connate or distinct?

5. GYNOECIUM: How many carpels make up this


syncarpous gynoecium and what is the placentation
type? The superior ovary of Dianthus sits upon a
ANDROGYNOPHORE (i.e. GYNOPHORE). What is the
difference between an ANDROGYNOPHORE and a
hypanthium?

Figure 7.10 Longitudinal section of a Dianthus flower.

6. PORTFOLIO DRAWING: Draw a floral diagram of a


longitudinal section (L.S.) of Dianthus as well as
a cross section (X.S.) of its ovary, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

FRUIT TYPE

▪▪ Fruit is a DENTICIDAL CAPSULE (a capsule with apical


teeth)

Silene spp. (Catchfly or Campion)


Agrostemma githago L. (Corn Cockle)

Most members of the Caryophyllaceae family, like Silene


species and A. githago, produce DENTICIDAL CAPSULES
with free central placentation and persistent connate
sepals (Figure 7.11). What type of gynoecium is required
to produce this fruit type (or any other type of capsule)?

Figure 7.11 Longitudinal section of an Agrostemma


githago denticidal capsule.

158 Integrative Biology 335: Plant Systematics


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Herbs with ANTHOCYANIN pigments


▪▪ Opposite, simple, entire leaves with connate, sheathing
leaf bases (creating SWOLLEN STEM NODES)

Lychnis chalcedonica L. (Maltesecross)

Review your vegetative terminology while observing some


of the major characteristics of Caryophyllaceae like the
swollen stem nodes (Figure 7.13).

Figure 7.12 Complex cyme from a Lychnis chalcedonica.


Complexity __________________________________

Arrangement __________________________________

Attachment __________________________________

Leaf venation __________________________________

Leaf apex __________________________________

Leaf base __________________________________

Leaf margin __________________________________

Figure 7.13 Opposite leaves and swollen nodes of a


Dianthus caryophyllus stem.

Lab 7 - Cactaceae, Caryophyllaceae, Portulacaceae and Polygonaceae 159


PORTULACACEAE
purslane family ♂ Ca2 Co5-∞ A5-∞ G 2-9

Eudicots: Caryophyllid Clade

FLORAL CHARACTERISTICS

▪▪ Two persistent sepals (may be bracts)


▪▪ Two to many ovules with basal or free central placentation

Portulaca grandiflora Hook. (Moss-rose Purslane)

To observe the floral morphology characteristic of the


Portulacaceae family you will examine a P. grandiflora
flower.

1. Looking at the provided photographs, what is the floral


symmetry?

2. CALYX: Identify the 2 persistent sepals (Figure 7.14).

Figure 7.14 Underside of a Portulaca grandiflora flower. 3. COROLLA: How many petals make up the corolla?

4. ANDROECIUM: Number of stamens (few or


numerous)?

5. GYNOECIUM: Note the one-loculed, partially inferior


ovary with many ovules (Figure 7.15). What is the
placentation type? Also, how many styles or style
branches are there? What does this tell you about the
type of gynoecium and the carpels that make it up?

6. Write out the floral formula for the P. grandiflora


flower.

Figure 7.15 Longitudinal section of a Portulaca


grandiflora flower.

160 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruits are capsules (sometimes CIRCUMSCISSILE


CAPSULES (PYXIS))

Portulaca oleracea L. (Common Purslane)

In P. oleracea, the fruit is a circumscissle capsule (PYXIS)


(Figure 7.16). In order for the seeds to dehisce, or be
released from the fruit, the top of the fruit comes off like a
lid.

Claytonia virginica L. (Spring Beauty)


Talinum teretifolium Pursh. (Quill Fameflower)
Figure 7.16 Pyxis of Portulaca oleracea.
Check out the other capsule types produced by C.
virginica and T. teretifolium (Figure 7.17).

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Produce BETALAIN pigments


▪▪ Leaves often succulent

Portulaca oleracea L. (Common Purslane)

The leaves of P. oleracea are succulent, a common


feature of Portulacaceae. Now take a minute to review
your vegetative terminology by drawing and labeling a leaf
in Box 3. Also fill in the blanks below.

Figure 7.17 Sequence from flower to capsule fruit in Complexity __________________________________


Talinum teretifolium.
Attachment __________________________________

Blade shape __________________________________


Box 3
Leaf apex __________________________________

Leaf base __________________________________

Leaf margin __________________________________

Lab 7 - Cactaceae, Caryophyllaceae, Portulacaceae and Polygonaceae 161


POLYGONACEAE
buckwheat family ♂ Ca3+3 (5) Co0 A3-9 G 2-3

Eudicots: Caryophyllid Clade

FLORAL CHARACTERISTICS

▪▪ Sepals often in 2 whorls of 3 (sometimes an inner sepal


fused with an outer one)
▪▪ Petals are absent

Polygonum capitatum Buch.-Ham. (Pinkhead


Smartweed)

Polygonum is Greek for “many-kneed” in reference to


the jointed stems characteristic of the genus. To observe
the floral morphology characteristic of the Polygonaceae
family you will examine a P. capitatum flower. Flowers
produced by Polygonaceae species are considered perfect
but incomplete. How is this possible?

Figure 7.18 Inflorescences of Polygonum coccineum.


1. Observe one of these tiny flowers. What is its
symmetry?

2. CALYX: Using a microscope, verify that there are 5


sepals making up the calyx. Typically the sepals for
Polygonaceae species will occur in 2 whorls of 3
(noted in the floral formula as 3 + 3). However, in P.
capitatum one of the inner sepals is fused with a sepal
in the outer whorl creating a total of 5 sepals. Also
notice that these sepals are white to pink in color - not
all sepals are green!

3. COROLLA: Petals, and thus the corolla, are absent in


members of the Polygonaceae Family.

4. ANDROECIUM: Note the arrangement of the 8 stamens


that make up the androecium.

5. GYNOECIUM: The ovary is trigonous (3-sided) and


has 3 capitate stigmas (Figure 7.19). Based on this,
how many carpels do you think fused to produce the
flower's syncarpous gynoecium? What is the ovary
position?
Figure 7.19 Inflorescences (Top) and flowers (Bottom) of
Polygonum capitatum.

162 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruits are triangular or lens-shaped achenes

Fagopyrum esculentum Moench (Buckwheat)


Rumex spp. (Docks)

Check out the triangular achenes produced by F.


esculentum and Rumex (Figures 7.20 and 7.21). What are
Figure 7.20 Achenes of Fagopyrum esculentum. some of the key features of an achene fruit? Also, how
many ovules were in the original flower to produce these
achenes?

Figure 7.21 Achenes of Rumex crispus.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Produce ANTHOCYANIN pigments


▪▪ Stipules are usually present and sheathing around stem,
called OCREA

Coccoloba uvifera L. (Seaside grape)


Polygonum pennsylvanicum L. (Pennsylvania
smartweed)

Find the sheathing stipules (OCREA) at the base of the


petioles surrounding the stem. This character is diagnostic
for Polygonaceae, and its shape and pubescence is often
used to identify species in the Polygonum genus. Please
label the OCREA, node, stem and petiole in Figure 7.22.

Figure 7.22 Stem of Polygonum pennsylvanicum


displaying its characteristic OCREA.

Lab 7 - Cactaceae, Caryophyllaceae, Portulacaceae and Polygonaceae 163


164 Integrative Biology 335: Plant Systematics
LABORATORY 8
CUCURBITACEAE, SALICACEAE, VIOLACEAE,
MALVACEAE, BRASSICACEAE AND ERICACEAE

PHYLOGENETIC RELATIONSHIPS

All of the families examined in today’s lab, with


the exception of Ericaceae, belong to the Rosid
clade.

Cucurbitaceae, along with begonias, comprise


the order Cucurbitales in the Fabids subclade.
Synapomorphies include inferior ovaries, strongly
intruded parietal placentae, unisexual flowers,
and the presence of cucurbitacins. The highly
modified androecium of Cucurbitaceae is similar
in appearance to the style and stigma, and
insects are fooled into visiting both staminate and
carpellate flowers. The family is very important
economically for its edible fruits (PEPOS).

Violaceae is included within the order


Malpighiales, an order in the Fabids subclade,
that is circumscribed on the basis of molecular
data only.

Salicaceae is also found in the order


Malpighiales. The monophyly of the family is supported primarily by molecular evidence, but also by the distinctive
salicoid teeth and associated gladular hairs, imperfect apetalous flowers and salicin. Though the flowers of the two
major genera in Salicaceae, Salix (willows) and Populus (i.e. poplar, cottonwood, aspen), may seem simple, they are
not primitive; they are actually highly reduced. Salix and Populus both provide lumber, wood pulp and ornamentals.
Additionally, the bark of Salix was used medicinally as a source of salicylic acid (the original Aspirin) to reduce
swelling and fever. Also included in Salicaceae are those genera previously treated in the “Flacourtiaceae”, such as
the ornamentals Oncoba, Casearia and Idesia.

Brassicaceae is found in the order Brassicales (15 families), and thus is part of the Malvids subclade.
Synapomorphies for Brassicaceae include sulfur-bearing glucosinolate compounds, the cruciform arrangement of the
petals and the unique structure of the endoplasmic reticulum. It includes many important food plants, such as capers,
radish, cabbage, kale, broccoli, cauliflower, brussels sprouts, turnips, Chinese cabbage, mustard and horseradish.
Canola oil is extracted from the seeds of Brassica napus. Arabidopsis is also incuded here, and is the most widely
used vascular plant in molecular biology!

Malvaceae, the Mallow family, is placed in the order Malvales along with eight other families. Phylogenetic analysis
of DNA sequences clearly place the order within the Malvid subclade; previously, many systematics had considered
Malvales to be related to the Urticales. Synapomorphies include mucilage (slime) canals and cavities, stellate hairs,
and connate sepals. Stamens are frequently numerous and develop centrifugally. Judd et al. place Theobroma cacao
(chocolate) within the Malvaceae family. Other economically important plants within the Malvaceae include Cola nitida
(cola seeds), Durio zibethinus (durian fruit), and Hibiscus esculentus (okra). You can have a wicked party serving just
these food items!

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 165


Lastly, Ericaceae, found in the order Ericales,
is distantly related to the other five families. It
is a member of the Asterid clade, a large and
specialized subgroup within the Eudicots. The
Ericales are a sister group to the Core Asterids.
Ericaceae is broadly circumscribed to include
five other families that are sometimes recognized
separately. The family is circumscribed as
monophyletic on the basis of phylogenetic
analysis of molecular data and includes many
showy ornamentals and plants with edible fruits
(e.g. blueberries, cranberries).

166 Integrative Biology 335: Plant Systematics


CUCURBITACEAE ♂ Ca5 Co⑤ A⑤ G0
cucumber family
♀ Ca5 Co⑤ A0
Eudicots: Rosid Clade
G③

FLORAL CHARACTERISTICS

▪▪ Imperfect flowers that are often yellow


▪▪ Staminate flowers: filaments and anthers connate and
anthers folded or bent

Cucumis spp. (Cucumbers and Melons)

To observe the floral morphology characteristic of the


Cucurbitaceae family you will examine and dissect
carpellate and staminate flowers of a Cucumis species.
Figure 8.1 A Citrullus vulgaris plant.
CARPELLATE FLOWER

1. Examine a carpellate flower. What is its symmetry?

2. CALYX and COROLLA: Determine the number of


sepals and petals. Is each perianth whorl connate or
distinct?

Male and female flowers of Citrullus


3. ANDROECIUM: Notice that the androecium is not
Figure 8.2
vulgaris, commonly known as watermelon. present in the carpellate flower. However, what
structure is present in place of the stamens?

Box 1

4. GYNOECIUM: How many stigmas and styles do you


see and what does this tell you about the number
of carpels fused to produce the flower’s syncarpous
gynoecium? Is the ovary position inferior or superior?
Make a cross-section (X.S.) of the ovary, observe the
single locule and then sketch the X.S. in Box 1. What
type of placentation does the ovary have?

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 167


STAMINATE FLOWER

1. Examine a staminate flower and notice its long


peduncle. What is its symmetry?

2. CALYX and COROLLA: Determine the number of


sepals and petals. Is each perianth whorl connate or
distinct?

Figure 8.3 Longitudinal sections of male and female


Citrullus vulgaris flowers.
3. ANDROECIUM: Count the number of connate stamens
that make up the androecium. Also note the folded
anthers. Why do you think the androecium of the male
flowers has evolved to look this way?

4. GYNOECIUM: Notice that the gynoecium is not


present in the male flower.T AND VEGETATIVE
CHARACTERISTICS

Figure 8.4 Male and female flowers of Ecballium


elaterium, or squirting cucumber.

FRUIT TYPE

▪▪ Fruit is a PEPO

Cucurbita moschata Duchesne. (Butternut Squash)


Cucumis sativus L. (Garden Cucumber)
Cucumis melo subsp. melo (Canteloupe)

Characteristic of this family, the ovaries of these


species mature into large, water-filled, thick-walled fruits
called PEPOS. In what ways does a PEPO differ from a
hesperidium and a berry?

Figure 8.5 Assortment of Cucurbitaceae flowers and


fruits.

168 Integrative Biology 335: Plant Systematics


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Herbaceous vines with TENDRILS


▪▪ Leaves palmately lobed or divided

Cucurbita spp. (Squashes and Pumpkins)

Review your vegetative terminology while observing the


two main features of Cucurbitaceae listed above on the
live plant specimen provided. The TENDRILS on the vine
allow it to climb (Figures 8.6 and 8.7)!

Figure 8.6 Herbaceous vine of Citrullus vulgaris.


Notice the male and female flowers. Complexity __________________________________

Arrangement __________________________________

Attachment __________________________________

Venation __________________________________

Figure 8.7 Cucurbita vine with pepo fruit.

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 169


VIOLACEAE
violet family ♂ CaZ5 CoZ5 A5 G③

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS

▪▪ Zygomorphic symmetry
▪▪ One petal has a NECTAR SPUR
▪▪ Two lower anthers have a nectary that protrudes into the
NECTAR SPUR
▪▪ Produce two types of flowers (CHASMOGAMOUS and
CLEISTOGAMOUS)

Viola spp. (Violets)

In the spring, most species produce normal petaliferous


flowers (CHASMOGAMOUS flowers) like you will see in lab
today, but in the summer they may also produce fertile
CLEISTOGAMOUS flowers.

1. Observe a flower (Figure 8.8). What is its symmetry?

2. CALYX: How many sepals make up the calyx? Are


they connate or distinct?
Figure 8.8 Viola tricolor flower exhibiting prominent
nectar guides.

3. COROLLA: How many petals make up the corolla and


are they connate or distinct? What is unusual about
the lowermost petal (Figure 8.9)?

4. ANDROECIUM: How many stamens are in the


androecium? Observe the small filaments, and the
anthers with their membranous appendages. The
lower two anthers each have a gland-like nectary
on them that extends into the spur of the lowermost
petal.

Figure 8.9 Longitudinal section of Viola with


characteristic NECTAR SPUR.

170 Integrative Biology 335: Plant Systematics


5. GYNOECIUM: What is the ovary position and what
type of insertion does the flower display? How many
carpels form the syncarpous gynoecium?

6. What are the small, unopened flowers located near


the base of a violet called (Figure 8.10)?

Figure 8.10 Viola sororia flowers.

FRUIT TYPE

▪▪ Fruits are loculicidal capsules that are explosively


dehiscent

Viola sororia Willd. (Common Blue Violet)


Viola pedata L. (Birdfoot Violet)

Check out the pictures of the loculicidal capsules produced


by V. sororia and V. pedata. Based on what you see, what
type of placentation did the original flower have?

Figure 8.11 Loculicidal capsules of Viola sororia.


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit is an herb

Viola sororia Willd. (Common Blue Violet)


Viola pedata L. (Birdfoot Violet)
Viola striata Aiton (Pansy)

Observe the herbarium specimens on display to see the


characteristic habit and some species diversity of the
Violaceae family (Figure 8.12). The State Flower of Illinois
is the “Blue Violet”. The only problem is that there are
several different species of blue violets native to Illinois,
and state law does not specify which one is the official
state flower.

Figure 8.12 Habit of Viola pedata.

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 171


SALICACEAE
♂ Ca0-x Co0 A2-∞ G0
willow family

Eudicots: Rosid Clade ♀ Ca 0-x Co0 A0 G 2-4

TWO MAJOR GENERA

▪▪ Populus spp. (Aspen, Poplar and Cottonwood)


▪▪ Salix spp. (Willows)

FLORAL CHARACTERISTICS

▪▪ Flowers imperfect and plant dioecious


▪▪ Calyx reduced to a disk in Populus or nectar glands in
Salix and often subtended by a bract
▪▪ Populus wind pollinated and Salix bee pollinated

Figure 8.13 Male catkin of Populus deltoides.

Populus deltoides Marsh. (Eastern Cottonwood)

The staminate flowers of Populus species exhibit a disk- or


cup-like calyx that the five to many dangling stamens are
attached to (Figures 8.13 and 8.14). The carpellate flowers
possess a similar calyx but instead have a syncarpous
gynoecium attached to it that is made up of 2 to 4 connate
carpels (Figure 8.15). Both staminate and carpellate
flowers are subtended by a fringed bract.

What type of inflorescence are both the staminate and


carpellate flowers clustered into?

What is the mechanism for pollination in Populus


Figure 8.14 Close-up of Populus deltoides male catkin. species? Why do you think this is the case based on the
morphology of the staminate and carpellate flowers?

Figure 8.15 Close-up of Populus deltoides female catkin.

172 Integrative Biology 335: Plant Systematics


Salix spp. (Willows)

In Salix species, the calyx of both staminate and carpellate


flowers are reduced to one or two basal nectary glands.
Evidence from vascular anatomy indicates that these
glands might represent reduced sepals (or even petals).
Typically, each staminate flower has only two stamens
while each carpellate flower has a syncarpous gynoecium
of 2 connate carpels and a superior ovary (Figure 8.16).
Both flowers are subtended by bracts which may be quite
pubescent as in S. discolor (pussy willow.)

Based on your class notes, the plant condition for


members of the Salicaceae family is DIOECY (DIOECIOUS).
What does this mean? Also, what is the mechanism of
pollination in Salix species? Why do you think this?

Figure 8.16 Individual male and female flowers of Salix


exigua, sandbar willow. Note that both sexes
of flowers have bracts and nectar glands.

FRUIT TYPE

▪▪ Fruits are capsules and contain hairy (COMOSE) seeds

Populus deltoides Marsh. (Eastern Cottonwood)


Salix interior Rowlee (Sandbar Willow)
Salix nigra Marsh. (Black Willow)

Check out the capsule fruits of P. deltoides, S. interior


and S. nigra and the COMOSE seeds emerging from them
(Figures 8.17 and 8.18). What do you suppose disperses
the seeds of these species?

Figure 8.17 Loculicidal capsules and comose seeds of


Populus deltoides.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit is a tree or shrub

Populus alba L. (White Poplar)


Salix spp. (Willow)

Examine the live material and herbarium specimens


on display to see sample vegetative material of the
Salicaceae family.

Figure 8.18 Loculicidal capsules of Salix exigua.

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 173


BRASSICACEAE (CRUCIFERAE)
mustard family
♂ Ca4 Co4 A4+2 G②

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS

▪▪ 4-merous flowers
▪▪ TETRADYNAMOUS stamens
▪▪ Ovaries are 2-locular with numerous parietal ovules

Brassica spp. (Canola or Rape)

To observe the floral morphology characteristic of the


Brassicaceae family you will dissect a Brassica flower.

1. Observe a flower (Figure 8.19). What is its symmetry?

Figure 8.19 Brassica flower with TETRADYNAMOUS


STAMENS.

2. CALYX and COROLLA: How many sepals and petals are


present? Do you detect any adnation or connation?
Note that the petals have an elongate claw and an
abruptly spreading blade.

3. ANDROECIUM: The androecium in this family is


described as TETRADYNAMOUS (Figure 8.20).
What does this term mean and how is this feature
accounted for in Brassicaceae’s floral formula?

Figure 8.20 Longitudinal section of Brassica flower.


4. GYNOECIUM: Observe the short, solitary style. The
ovary is bicarpellate and is divided into two locules
by a thin septum. The ovules are borne in 2 rows and
Box 2 are separated by this partition. As the ovary matures,
the septum forms a persistent REPLUM. Make a cross
section of the ovary and then draw and label it in Box
2.

5. DRAWING PORTFOLIO: Draw a floral diagram of a


longitudinal section (L.S.) of Brassica as well as
a cross section (X.S.) of its ovary, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

174 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruits are SILIQUES or SILICLES with a REPLUM


▪▪ Seeds contain a bent embryo

Hesperis matronalis L. (Dame’s Rocket)


Lunaria spp. (Money Plant)
Thlaspi arvense L. (Field Pennycress)

Although Brassicaceae flowers are diagnostic for the


family, their uniformity can cause many taxonomic
problems; taxonomic distinction among taxa largely rests
on characters of the mature fruit. If fruits are not available,
one can make an educated guess about what kind of fruit
will be formed (SILIQUE or SILICLE) by examining the ovary
of the flower. A SILIQUE forms from a gynoecium with a
long and thin ovary, whereas a SILICLE forms from a short
and broad ovary (Figure 8.21). Thus, what do you think is
the major difference between a SILIQUE and a SILICLE?

When a fruit is ready to disperse its seeds, the two valves


Figure 8.21 Silique fruits of Hesperis matronalis and fall away leaving the persistent REPLUM. Observe the
silicle fruits of Thlaspi species. Lunaria specimen on display - only the REPLUMS are left!
In this case the valves have already fallen off, thereby
releasing the seeds.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Usually annual, biennial or perennial herbs

Arabidopsis thaliana (L.) Heynh. (Mouseear Cress)

Since the 1900s this herb, A. thaliana, has been used as a


model organism by plant biologists. Aside from being easy
to propagate, its small size and rapid life cycle (six weeks
from germination to fruiting) make A. thaliana an ideal
candidate for manipulative experiments. Additionally, in
2000 this species became the first plant to have its entire
genome sequenced.

Figure 8.22 Raceme of Brassica rapa, or canola.

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 175


MALVACEAE
mallow family A ∞
♂ Ca⑤ Co5 G 5-∞

Eudicots: Rosid Clade

FLORAL CHARACTERISTICS

▪▪ Sepals often subtended by bracts (EPICALYX)


▪▪ Petals adnate to staminal tube
▪▪ MONADELPHOUS stamens with unilocular and crescent-
shaped anthers

Hibiscus rosa-sinensis L. (Rose-of-China)

To observe the floral morphology characteristic of the


Malvaceae family you will examine and dissect a H. rosa-
sinensis flower.

1. Observe a flower (Figure 8.23). What is its symmetry?


Figure 8.23 A Hibiscus flower with its characteristic
MONADELPHOUS staminal tube.

2. CALYX: Distinguish between the EPICALYX (an


involucre or whorl of bracts immediately subtending
the calyx) and the connate calyx (Figure 8.24). How
many sepals make up the calyx?

3. COROLLA: The petals are distinct and are adnate to


the base of the androecium’s staminal tube. Notice
how the floral formula is written to account for the
adnation of these two parts. How many petals are
there?

4. ANDROECIUM: Observe the MONADELPHOUS (in Greek:


“mon” = one and “adelphous” = brother) stamens
characteristic of this family. This term refers to the
connation of the filaments at their base forming a
Figure 8.24 Longitudinal section of a Hibiscus flower.
Notice the EPICALYX subtending the sepals. single staminal tube. At the top of the staminal tube
the filaments branch off, each bearing a crescent-
shaped anther.

176 Integrative Biology 335: Plant Systematics


5. GYNOECIUM: How many carpels have fused to
produce this syncarpous gynoecium? (Hint: Count the
stigmas or upper style branches.) What is the ovary
position and the placentation type?

6. DRAWING PORTFOLIO: Draw a floral diagram of a


longitudinal section (L.S.) of H. rosa-sinensis as well
as a cross section (X.S.) of its ovary, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

Abutilon spp. (Abutilon or Flowering Maple)

If available, dissect one of the flowers of Abutilon. You will


notice that unlike H. rosa-sinensis, there is no EPICALYX
present and the connate carpels each contain two or more
ovules.

Figure 8.25 Capsule fruit of Iliamna remota, commonly


called Kankakee mallow.

FRUIT TYPE

▪▪ Fruits are schizocarps or capsules

Iliamna remota Greene (Kankakee Mallow)


Abutilon theophrastii Medik. (Velvetleaf)

The fruit type produced by both these species is a


capsule, a dry dehiscent fruit type that will open to release
multiple seeds (Figure 8.25).

Gossypium spp. (Cotton)

Observe the dehiscing capsule fruit of Gossypium. The


long hair on the seeds is the cotton fiber that we all know
and love (Figure 8.26). What is the term used to describe
the “hairiness” of the seeds?

Figure 8.26 Capsule fruit of Gossypium, or cotton.


Notice the long hairs attached to the seeds.

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 177


Box 3 HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Leaf and stem surfaces commonly have STELLATE HAIRS


▪▪ Palmately lobed or veined leaves
▪▪ Plants often contain mucilaginous sap

Abutilon spp. (Abutilon or Flowering Maple)

Observe a piece of this plant’s leaves under a microscope


to see the STELLATE HAIRS on its surface. How would you
describe the leaf margin and leaf venation of Abutilon?
Sketch out a typical Malvaceae leaf, including its stellate
hairs, in Box 3.

Althaea officinalis L. (Marsh Mallow)

The first marshmallows were originally made during the


early 19th century in France by whipping the mucilaginous
sap of this plant’s roots and sweetening it (Figure
8.27). By the late 19th century the mucilaginous sap in
marshmallows was replaced with gelatin mixed with a
modified corn starch. Yum!

Figure 8.27 Marshmallows, yum!

178 Integrative Biology 335: Plant Systematics


ERICACEAE ♂ Ca5 Co⑤ A10 G 2-10

heather family
OR
♂ Ca5 Co⑤ A10

Eudicots: Asterid Clade
G⑤

FLORAL CHARACTERISTICS

▪▪ Corolla often URCEOLATE or CAMPANULATE


▪▪ Anthers often APPENDAGED and opening by pores with
pollen in tetrads
▪▪ Gynoecium exhibits axile placentation and a hollow style

Rhododendron spp. (Rhododendron or Azaleas)

To observe the floral morphology characteristic of


the Ericaceae family you will examine and dissect a
Rhododendron flower.

1. Observe a flower (Figure 8.28). What is its symmetry?


Is this typical in the Ericaceae family?

2. CALYX: How many sepals make up the calyx? Are


Figure 8.28 Longitudinal section of a Rhododendron they connate or distinct?
‘PJM’ flower.

3. COROLLA: The CAMPANULATE shaped corolla of this


flower is formed through the connation of its five
petals.

4. ANDROECIUM: How many stamens make up the


androecium? Locate and examine the pubescent,
elongate filaments and the short anthers with terminal
pores which are characteristic of Ericaceae species.

5. GYNOECIUM: Rhododendron’s syncarpous gynoecium


is composed of 5 carpels, and, like most genera
of Ericaceae, the ovary is superior. Make a cross-
section (X.S.) of the ovary (Figure 8.29). What is the
placentation type?

Figure 8.29 Cross section of a Rhododendron ‘PJM’


flower’s base.

Lab 8 - Cucurbitaceae, Violaceae, Salicaceae, Brassicaceae, Malvaceae and Ericaceae 179


Pieris japonica (Thunb.) D. Don ex G. Don. (Japanese
Andromeda)

This 5-merous flower has 5 distinct sepals, an ovoid-


URCEOLATE corolla of 5 connate petals, 10 stamens
with anthers having a pair of awns near the base and a
5-locular ovary containing numerous ovules (Figure 8.30).

Figure 8.30 Urceolate flowers of Pieris japonica.

FRUIT TYPE

▪▪ Fruit can be capsules, berries or drupes

Vaccinium corymbosum L. (N. Highbush Blueberry)

The fruit produced by V. corymbosum, a blueberry, is also


botanically considered a berry (Figures 8.31 and 8.32). Is
it produced from a superior or an inferior ovary? How can
you tell?

HABIT AND VEGETATIVE CHARACTERISTICS


Figure 8.31 Longitudinal section of Vaccinium
corymbosum flower.
▪▪ Mainly shrubs that prefer acidic soil
▪▪ Leaves simple, mostly entire and often thick, leathery and
evergreen

Kalmia angustifolia L. (Sheep Laurel)


Rhododendron groenlandicum (Oeder) Kron & Judd
(Bog Labrador Tea)

Observe the herbarium specimens on display to see


the few vegetative characteristics representative of the
Ericaceae family.

Figure 8.32 Berry fruit of Vaccinium corymbosum.

180 Integrative Biology 335: Plant Systematics


LABORATORY 9
FABACEAE, ONAGRACEAE AND EUPHORBIACEAE

PHYLOGENETIC RELATIONSHIPS

All of the families we will examine today belong to


the Rosid clade.

Fabaceae (or Leguminosae) is a member of the


order Fabales in the Fabids subclade. This family
is allied closely with the Rosales (e.g. Rosaceae,
Moraceae), Cucurbitales (e.g. Cucurbitaceae)
and Fagales (e.g. Betulaceae, Fagaceae). The
monophyly of Fabales is supported by molecular
data, vessel elements with single perforations, a
large green embryo, and the presence of ellagic
acid. Fabaceae is the third largest family in the
world, with some 18,000 species. It is also of
incredible economic importance, second only
to the grass family (Poaceae). The Fabaceae
contain beans, soybeans, lentils, peanuts,
chickpeas and a host of other agricultural and
ornamental species, such as alfalfa, clover, vetch,
orchid tree, redbud, lupine, sensitive plant and
acacia. The family is typically divided into three
subfamilies (Mimosoideae, Caesalpinioideae
and Faboideae), although some taxonomists,
such as Cronquist, have considered these taxa as three separate families. Recent analyses have shown that the
Caesalpinioideae are paraphyletic, with some members more closely related to Mimosoideae and others more closely
related to Faboideae than they are to each other. The redbuds (Cercis spp.) may be sister to the rest of the family.

Onagraceae belongs within the order Myrtales of the Rosid clade. However, the taxonomic placement of this order
is uncertain; in the phylogeny it falls outside of the Fabids and Malvids subclades. The monophyly of Onagraceae
is supported by having pollen grains associated with VISCIN THREADS, flowers with a well-developed hypanthium,
4-merous flowers, 4 fused carpels and an inferior ovary. Fuchsia, Oenothera and Clarkia are ornamentals with showy
flowers.

Euphorbiaceae belongs in the order Malpighiales alongside the Salicaceae and Violaceae in the Fabid clade. It is a
large and extremely diverse family, with some 300 genera and over 6,000 species. Many genera produce chemicals
that are medicinally as well as industrially important, such as Hevea brasiliensis (rubber plant) and Ricinus communis
(caster oil). Some other members are eaten (e.g. Manihot esculenta (cassava)). The family also occupies a wide
range of habitats, from wet tropical rain forests to desert environments where species have evolved a remarkable
resemblance to the Cactaceae. Within the Euphorbiaceae is the large tribe Euphorbieae (containing the genus
Euphorbia, or poinsettia), which is monophyletic on the basis of their inflorescence type (CYATHIUM, pl. CYATHIA).

Lab 9 - Fabaceae, Onagraceae and Euphorbiaceae 181


FABACEAE (LEGUMINOSAE)
legume family ♂ Ca5 Co5 A5-∞ G1

Eudicots: Rosid Clade

THREE SUBFAMILIES

▪▪ MIMOSOIDEAE
▪▪ CAESALPINIOIDEAE
▪▪ FABOIDEAE

Fabaceae is divided into three distinct subfamilies. The


subfamilies can be differentiated based on floral and
vegetative characteristics.L

FLORAL CHARACTERISTICS

▪▪ Perfect flowers
▪▪ Single, superior carpel with marginal placentation

Though all members of Fabaceae have perfect flowers


containing a single, superior carpel with marginal
placentation, the 3 subfamilies differ in their floral
Figure 9.1 Legumes of Phaseolus vulgaris. symmetry, corolla arrangement and androecia.

Floral Corolla
Subfamily Floral Formula Androecium
Symmetry Arrangement

Petals connate - 10 or more stamens,


MIMOSOIDEAE ♂ Ca⑤ Co⑤ A 10-∞ G1
♀ Actinomorphic and forming a and forming a tube
(Mimosa Subfamily)
tube - Pollen in POLLINIA

BANNER petal - 10 (or fewer) stamens


CAESALPINIOIDEAE
(Caesalpinia Subfamily)
♂ Ca5 CoZ5 A10 G1
♀ Zygomorphic internal to - Anthers often
lateral petals opening by pores

BANNER petal
outermost, 2
10 stamens either...
distinct lateral
FABOIDEAE
(Bean / Pea Subfamily)
♂ Ca 5 CoZ5 A 9 +1 or 10 G1
♀ Zygomorphic WING petals,
- DIADELPHOUS
- MONADELPHOUS
and 2 petals
- Distinct
fused to form
the KEEL

182 Integrative Biology 335: Plant Systematics


MIMOSOIDEAE
Calliandra spp. (Powderpuff Tree)

The name of this genus is from the Greek, “kalos”


(beautiful) and “andros” (stamens), referring to its
conspicuous thin, red stamens. The key to remembering
this subfamily is the phrase, “TUBE IN A TUBE IN A TUBE”,
referring to the connation of the sepals, the petals and (at
least the base of) the stamens (Figure 9.2).

1. Observe a flower. What is its symmetry?

2. CALYX: Notice that the sepals are fused into a tube, or


connate. How many are present?

Figure 9.2 An Inga flower.

3. COROLLA: The petals are also connate forming a tube.


How many are there?

4. ANDROECIUM: These flowers contain 10 to many


stamens that are fused at the base, but have distinct
filaments (Figure 9.3). As mentioned above, the genus
and common name of these flowers refer to the long,
thin stamens that project far above the corolla tube.

5. GYNOECIUM: Examine the monocarpous gynoecium.


Now sketch out a longitudinal section of the flower in
Box 1.

Figure 9.3 A longitudinal section of an Inga flower.

Box 1

Lab 9 - Fabaceae, Onagraceae and Euphorbiaceae 183


CAESALPINIOIDEAE
Caesalpinia pulcherrima (L.) Sw. (Pride-of-Barbados)

To observe the floral morphology characteristic of


the Fabaceae subfamily Caesalpinioideae, you will
examine and dissect a C. pulcherrima flower. The genus
Caesalpinia was named for Andreas Caesalpini (1519 -
1603), an Italian botanist and physician.

1. Observe a flower (Figure 9.4). What is its symmetry?

2. CALYX: How many sepals are present? Do you detect


Figure 9.4 A Caesalpinia pulcherrima flower.
any adnation or connation?

Box 2

3. COROLLA: Observe the five petals in this flower. There


are 2 lateral petals (the wings), a BANNER petal that
is internal to the lateral petals, and two lower petals
that are distinct from one another. The position of the
BANNER petal is critical!

4. ANDROECIUM: How many stamens make up the


androecium in this flower? The anthers in Fabaceae
subfamily Caesalpiniodeae often open by pores to
release pollen.

5. GYNOECIUM: Examine the single carpel with its


superior ovary. What is the gynoecium type and
insertion type found in this flower? Sketch out a
longitudinal section of the flower in Box 2.

Cercis canadensis L. (Eastern Redbud)

These trees bloom very early in the spring prior to


their leaves flushing out. What is the position of the
inflorescences based on the photo/material provided?

Though the species belongs to Caesalpinioideae, C.


canadensis flowers have connate sepals (not shown
in the floral formula for the subfamily) and “pea-like” or
Faboideae subfamily-like petals (i.e. 2 WING petals, 2 KEEL
petals and 1 BANNER petal which is inside the other petals)
(Figure 9.5).
Figure 9.5 Cercis canadensis: a flower (top) and a
longitudinal section of a flower (bottom).

184 Integrative Biology 335: Plant Systematics


FABOIDEAE
Lathyrus odoratus L. (Sweetpea)

To observe the floral morphology characteristic of the


Fabaceae subfamily Faboideae you will examine and
dissect a L. odoratus flower. The genus name Lathyrus is
from the Greek meaning “pea”.

1. Observe a flower (Figure 9.6). What is its symmetry?

Figure 9.6 Flowers of Lathyrus latifolius.

2. CALYX: How many sepals are present? Do you detect


any adnation or connation?

3. COROLLA: Observe the five petals in this flower. There


are 2 lateral petals (the WINGS), a BANNER petal that is
outermost to the lateral petals, and 2 lower petals that
are fused to form the KEEL.

4. ANDROECIUM: There are 10 stamens in this flower,


9 of which are connate and 1 is distinct (Figure 9.7).
What term can be used to describe them?

5. GYNOECIUM: Examine the carpel of this monocarpous


Figure 9.7 Longitudinal section of a Lathyrus latifolius gynoecium. What is the ovary position? Sketch out a
flower. Notice the 9 connate stamens and 1 longitudinal section of the flower in Box 3.
distinct stamen.

Box 3

Lab 9 - Fabaceae, Onagraceae and Euphorbiaceae 185


FRUIT TYPE

▪▪ Fruit is a LEGUME

There is enormous variation in the morphology of


the Fabaceae’s LEGUME fruits. Observe the LEGUMES
produced by species in the 3 subfamilies: Mimosoideae,
Caesalpinioideae and Faboideae.

Examples

MIMOSOIDEAE
Figure 9.8 Legumes of Gymnocladus dioicus.
Albizia julibrissin Durazz. (Mimosa Tree)

CAESALPINIOIDEAE
Cercis canadensis L. (Eastern Redbud)
Gleditsia triacanthos L. (Honey Locust)
Gymnocladus dioicus (L.) K. Koch (Kentucky Coffeetree)

FABOIDEAE
Baptisia alba (L.) Vent. var. macrophylla (Larisey) Isely
(Largeleaf Wild Indigo)
Desmodium illinoense A. Gray (Illinois Ticktrefoil)
Glycine max (L.) Merr. (Soybean)
Phaseolus vulgaris L. (Common Bean)
Pisum sativum var. saccharatum L. (Snow Pea)
Figure 9.9 Legumes of Glycine max.

Box 4 Phaseolus vulgaris L. (Common Bean)


Pisum sativum var. saccharatum L. (Snow Pea)

Based on what you see in these examples, what was the


gynoecium type and the placentation type of the original
flower? How many sutures does a mature LEGUME dehisce
along?

Dissect a bean or snow pea and sketch it in Box 4. Label


all of the fruit or remnant flower structures listed in the
table below.

Flower Part Fruit Part


Pedicel ---
Sepals ---
Ovary wall Pericarp
Ovule Seed

186 Integrative Biology 335: Plant Systematics


LEGUME: Cercis canadensis L. (Fabaceae)
FOLLICLE: Aquilegia canadensis L. (Ranunculaceae)
SILIQUE: Hesperis matronalis L. (Brassicaceae)

What are the major differences among LEGUMES,


FOLLICLES and SILIQUES? (HINT: Think about the original
gynoecium type and the number of sutures the fruit has.)

Figure 9.10 The seeds of many legumes are beans!

SEED MORPHOLOGY

▪▪ Seeds lack an endosperm

Check out the seeds from the three Fabaceae subfamilies.


Make sure that you can tell if there is a PLEUROGRAM and,
if there is, describe its shape (Figures 9.11 and 9.12).

Figure 9.11 U-shaped pleurogram of Albizia julibrissin.

Examples

MIMOSOIDEAE: U-shaped groove

Albizia julibrissin Durazz. (Mimosa Tree)

CAESALPINIOIDEAE: O-shaped groove

Gleditsia triacanthos L. (Honey Locust)

Figure 9.12 O-shaped, or closed, pleurogram of a FABOIDEAE: No PLEUROGRAM


caesalpinoid legume.
Phaseolus vulgaris L. (Common Bean)

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Most species have root nodules containing nitrogen-fixing


bacteria

Examine the vegetative characters displayed by members


of the three Fabaceae subfamilies (Figure 9.13). Make
sure that you can identify a PULVINUS (a cushion-
Figure 9.13 Pinnately compound leaves of Mimosa like swelling at the base of a leaf or leaflet stalk) and
pudica. determine leaf complexity (i.e. pinnately compound,
bipinnately compound or palmately compound).

Lab 9 - Fabaceae, Onagraceae and Euphorbiaceae 187


Subfamily Habit Leaf Complexity PULVINUS Examples

Mostly tropical
Bipinnately Albizia julibrissin Durazz. (Mimosa Tree)
MIMOSOIDEAE and subtropical
compound
Present
Mimosa pudica L. (Sensitive Plant)
trees

Mostly tropical Pinnately or


Chamaecrista fasciculata (Michx.) Greene
CAESALPINIOIDEAE and subtropical bipinnately Absent
Gleditsia triacanthos L. (Honey Locust)
trees compound

Usually pinnately Amorpha canescens Pursh (Leadplant)


Mostly herbs,
compound, Lupinus spp. (Lupine)
FABOIDEAE some trees and
sometimes palmately
Absent
Phaseolus spp. (Beans)
shrubs
compound Trifolium pretense L. (Red Clover)

Mimosa pudica L. (Sensitive Plant)

Touch a leaflet of the sensitive plant! The species name,


“pudica” is very appropriate for this plant in that it means
bashful. Touching the leaves of M. pudica stimulates it to
release chemicals, thereby causing the redistribution of
water in the cells. The unequal distribution of water is what
allows the leaves to fold up (Figure 9.14). This mechanism
is speculated to be a defense against herbivory. Also, take
a minute to examine the PULVINUS of M. pudica leaves.

Complexity __________________________________

Attachment __________________________________

Figure 9.14 Pinnately compound leaves of Mimosa


pudica displaying a PULVINUS.

188 Integrative Biology 335: Plant Systematics


ONAGRACEAE
evening primrose family ♂ Ca4 Co4 A8

Eudicots Rosid Clade G④

INFLORESCENCE TYPE

Gaura mollis James (Velvetweed)


Oenothera biennis L. (Common Evening Primrose)

Examine the herbarium specimens provided. What is the


inflorescence type of each?

HABIT AND VEGETATIVE CHARACTERISTICS

FLORAL CHARACTERISTICS

▪▪ 4-merous flowers with a HYPANTHIUM and an inferior ovary


▪▪ Pollen sometimes with VISCIN THREADS
Figure 9.15 Flower of a cultivated Fuchsia.

Fuchsia triphylla L. (Fuchsia)

To observe the floral morphology characteristic of the


Onagraceae family, you will examine and dissect a F.
triphylla flower.

1. Observe the flowers (Figure 9.15). What symmetry


type do they display?

2. CALYX, COROLLA and ANDROECIUM: How many


sepals, petals and stamens are present in this flower?
The adnation of these three floral series forms what
structure (Figure 9.16)?

3. GYNOECIUM: How many carpels make up the


gynoecium in this flower? Are they connate or
distinct? Is the ovary position superior or inferior?
What is the insertion type?
Figure 9.16 Longitudinal section of a Fuchsia flower.

Lab 9 - Fabaceae, Onagraceae and Euphorbiaceae 189


4. DRAWING PORTFOLIO: Draw a floral diagram of a
longitudinal section (L.S.) of F. triphylla, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

FRUIT TYPE

▪▪ Fruit is a capsule or berry

Oenothera speciosa Nutt. (Pinkladies)

Like Fuchsia triphylla, Oenothera speciosa has a


HYPANTHIUM (floral cup) (Figure 9.17). Characteristic
of Onagraceae, this species also has VISCIN THREADS
Figure 9.17 Longitudinal section of an Oenothera connecting its pollen grains (Figure 9.18).
speciosa flower.

Examine the Oenothera fruit on display (Figure 9.19).


What is the fruit type (be specific)? What type of
gynoecium is required to produce this fruit type?

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Herbs or shrubs
Oenothera flower characteristics. ▪▪ No stipules
Figure 9.18

Clarkia spp. (Clarkia)

The genus Clarkia has more than 30 species ranging


from western North America to northern South America.
The genus is named for William Clark (1770-1838), the
co-leader of the Lewis and Clark expedition. Notice the
herbaceous habit of the plant.

Figure 9.19 Fruit produced by Oenothera biennis.

190 Integrative Biology 335: Plant Systematics


EUPHORBIACEAE
♂ Ca0 Co0 A1 G0
spurge family

Eudicots: Rosid Clade ♀ Ca 0 Co0 A0 G③

FLORAL CHARACTERISTICS

▪▪ Imperfect flowers
▪▪ Typically has a CYATHIUM inflorescence with nectar glands
and petal-like appendages
▪▪ 3 connate carpels make up the syncarpous gynoecium
▪▪ Superior ovary is 3-locular with 1 apical ovule per locule

Three Euphorbia cotonifolia cyathia at


There are two types of flowers in the Euphorbiaceae
Figure 9.20
different stages of female flower maturity. family: the “Euphorbia” type (restricted to the genus
Euphorbia) and the “Normal” type (like Jatropha species).
In this class we will focus on the “Euphorbia” type flowers.
Within the entire family though, flowers are imperfect
and the plants are monoecious or dioecious. What is the
difference between the monoecious and dioecious plant
condition?

Euphorbia cotinifolia L. (Mexican Shrubby Spurge)


Euphorbia corollata L. (Flowering Spurge)
Euphorbia millii Desmoul. (Crown-of-Thorns)
Euphorbia fulgens Karw. (Scarlet-Plume)
Figure 9.21 Euphorbia corollata CYATHIA.
In the Euphorbia genus the flowers are highly reduced and
what you see is not a “flower”, but actually a specialized
inflorescence called a CYATHIUM (pl. CYATHIA; Figures 9.20
and 9.21). The petal-like structures (appendages) are
actually glands that secrete nectar. Embedded within the
cup-like CYATHIUM are the staminate flowers which are
each reduced to a single stamen (Figure 9.22). The single
carpellate flower arises from the center of the cyathium on
a stalk and is comprised of three carpels, each with one
ovule. The perianth parts are absent. Think of a CYATHIUM
as a highly reduced cymose inflorescence with associated
bracts.

DRAWING PORTFOLIO: Draw a floral diagram of an E.


fulgens CYATHIUM, making sure to include the floral formula
for the carpellate and staminate flowers. Follow the format
as instructed in Lab 3: Floral Terminology.GETATIVE
CHARACTERISTICS

Figure 9.22 Longitudinal section of an Euphorbia milii


CYATHIUM.

Lab 9 - Fabaceae, Onagraceae and Euphorbiaceae 191


FRUIT TYPE

▪▪ Fruit is a SCHIZOCARP splitting into MERICARPS

Euphorbia corollata L. (Flowering Spurge)

With what you know about the gynoecium of Euphorbia


female flowers, how many MERICARPS do you think the
SCHIZOCARP of this species will split into (Figure 9.23)?

Figure 9.23 Schizocarp fruits of Euphorbia corollata.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Succulent plants often with milky sap

Euphorbia millii Desmoul. (Crown-of-Thorns)

According to legend, E. millii was the species that was


used to make the crown of thorns worn by Jesus during
the Crucifixion, hence its common name (Figure 9.24).
Examine the plant’s succulent leaves and the slender
SPINES that cover its stem. Now take a minute to review
your vegetative terminology by filling in the blanks below.

Complexity __________________________________
Figure 9.24 Vegetative features and CYATHIA of
Euphorbia millii. Attachment __________________________________

Blade shape __________________________________

Leaf apex __________________________________

Leaf margin __________________________________

Euphorbia spp.

Notice how these Euphorbia species resemble members


of the Cactaceae family with their succulent stems (Figure
9.25). What feature do they have though that Cactaceae
species do not?

Figure 9.25 Milky sap is common in Euphorbiaceae


succulents.

192 Integrative Biology 335: Plant Systematics


LABORATORY 10
ASCLEPIADACEAE, SOLANACEAE, OLEACEAE AND
CAPRIFOLIACEAE

PHYLOGENETIC RELATIONSHIPS

All of the families we will examine today belong to


the Asterid clade.

The Asterid clade is a large and specialized


subgroup of the angiosperm eudicot (tricolpate)
clade. It is hypothesized to be monophyletic
on the basis of molecular evidence, as well
as on the shared presence of ovules having
only a single integument and a thin-walled
megasporangium. The “Core Asterid” clade
is supported as monophyletic on the basis of
the number of stamens equaling the number
of petals, epipetalous stamens, an obviously
sympetalous corolla, and molecular data. This
group approximates the subclass Asteridae of
Cronquist.

The family Apocynaceae, as circumscribed on the


basis of cladistic analyses of both morphological
and molecular data, is paraphyletic. The
family Asclepiadaceae (treated as subfamily
Asclepiadoideae) arises from within Apocynaceae.
In this class, for the sake of convenience only, we will continue to treat the milkweeds in their own family,
Asclepiadaceae. This family is placed in the order Gentianales of the Lamiids subclade.

Solanaceae and Oleaceae belong to the orders Solanales and Lamiales, respectively, also in the Lamiids subclade.

Caprifoliaceae (and Adoxaceae) are members of the order Dipsacales, of the Campanulids subclade. Traditionally
these families were combined as one (Caprifoliaceae), but recent analyses treat them as sister families.
Caprifoliaceae is characterized by opposite leaves, a zygomorphic corolla (with epipetalous stamens), and an inferior
ovary. The showy flowers of Caprifoliaceae are pollinated by various nectar-gathering insects (primarily bees and
wasps) and birds.

Lab 10 - Asclepiadaceae, Solanaceae, Oleaceae and Caprifoliaceae 193


ASCLEPIADACEAE
milkweed family A⑤
♂ Ca⑤ Co⑤ G 2

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS

▪▪ Petals often a CORONA (consisting of a HORN & HOOD)


▪▪ Filaments, anthers and stigma fused into a GYNOSTEGIUM
▪▪ Pollen in pollinia and pollinia of adjacent anther
sacs connected by TRANSLATOR ARMS with gland or
CORPUSCULUM
▪▪ Carpels connate above and distinct below

Asclepias curassavica L. (Bloodflower)

To observe the floral morphology characteristic of


Asclepiadaceae, you will examine and dissect an
Asclepias curassavica flower. Make sure to look at the
provided diagrams and read through the verbal description
of the floral parts that follow to understand the morphology
Figure 10.1 An umbel inflorescence of Asclepias of this complicated flower.
curassavica.

1. Observe a flower. What is its symmetry?

1. CALYX: The green calyx is hidden under the reflexed


corolla (it’s easier to see the calyx on an unopened
flower). Are the sepals fused? (Typically they are in
the family, but in this species they are not.)

2. COROLLA: Arising from the red corolla is a yellow


CORONA (Figure 10.2). This CORONA consists of a
petaloid HOOD and a HORN (also called the BEAK). The
beak is exserted from the HOOD (meaning it sticks out)
and at its base (that is, within the HOOD) are nectaries.
The lower, inner part of the CORONA is attached to the
GYNOSTEGIUM.

3. ANDROECIUM and GYNOECIUM: Between each


CORONA element (HORN + HOOD) is a longitudinal slit
called the stigmatic slit. At the top of the slit is a dark
brown structure called a gland or CORPUSCULUM.
At the base of the slit is the receptive portion of the
Figure 10.2 Close-up of an Asclepias curassavica flower. stigma (you can’t see this). If you take a sharp probe
or needle and have a steady hand, you can run your
probe up the slit to the base of the gland. This gland

194 Integrative Biology 335: Plant Systematics


is connected to two TRANSLATOR ARMS which in turn
each connect to a pollen sac called a POLLINIUM (pl.
POLLINIA). If you lift the gland up and away from the
flower, you can pull out the two pollinia. Each of these
pollinia actually belong to adjacent anthers as the
anthers are positioned opposite the CORONA elements
and are alternate with the petals. The pollen within
each POLLINIUM is united into a large, waxy mass. The
two carpels have fused to produce the gynoecium in
this flower and they are united at their summit into
a large stigmatic head. The anthers are united, and
thus adnate to the stigmatic head. The large region
in the center of the flower (formed from the fusion of
the stigma to the anthers) is called the GYNOSTEGIUM.
Make a longitudinal cut through the flower and locate
the two ovaries (Figure 10.3). Within each ovary are
marginal ovules.

4. Label all the parts of the Asclepias flower on the line


drawing in Figure 10.4, using the live specimen and
the photographs provided.
Figure 10.3 Longitudinal section of an Asclepias
curassavica flower.

Stapelia spp. (Carrion Flowers)

Check out the photos of these unique plants. As their


common name suggests, the flower blossoms smell like
rotting meat to attract their major pollinators, flies.L

POLLINATION MECHANISM

Asclepiadaceae species have evolved a unique


mechanism for pollination. Because the apex of the
GYNOSTEGIUM is polished and slippery, an insect’s leg will
tend to slide off it and into the STIGMATIC SLIT between
two adjacent anthers. When the leg is pulled back up
it may be caught by the wishbone-shaped translators
and CORPUSCULUM. If the insect is strong, such as a
bumblebee, the pollinia are pulled free from the anthers
but remain attached to the insect’s leg as it flies to another
flower. (Weaker insects may be unable to loosen the
pollinia and are held so firmly that they die on the flower.)
At one of the subsequent flowers that is visited, the leg will
again slip off the top of the GYNOSTEGIUM and down into
the STIGMATIC SLIT. But this time, as the leg is pulled back
there is the possibility that part or all of one pollinium may
become lodged in the slit and break loose from the insect’s
foot. With the pollinium now lodged in the groove of a
different flower, it is adjacent to the receptive area of the
stigma and cross-pollination is achieved. Fertilization will
follow after the pollen grains in the waxy mass germinate
and their pollen tubes grow through the stigma, down the
styles, into the ovaries, into the ovules and enter into the
Figure 10.4 Diagram of an Asclepias flower. embryo sac. And voila, more milkweeds!

Lab 10 - Asclepiadaceae, Solanaceae, Oleaceae and Caprifoliaceae 195


FRUIT TYPE

▪▪ Fruit is a follicle containing seeds with COMA of hairs

Asclepias incarnata L. (Swamp Milkweed)


Asclepias syriaca L. (Common Milkweed)

Observe the milkweed fruits (follicles) provided (Figure


10.5). What are the key characteristics of the follicle fruit
Figure 10.5 An Asclepias syriaca flower with follicles. type? Based on the tufts of hair (COMA) on the seeds, what
is the seed dispersal mechanism (Figure 10.6)?

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Plants contain milky sap


▪▪ Leaves are simple with entire margin and opposite
arrangement

Asclepias sullivantii Engelm. ex A. Gray (Prairie


Milkweed)
Asclepias tuberosa L. (Butterfly Milkweed)

Check out the provided herbarium specimens for these


two Asclepias species. Both species contain milky sap
and can be found in higher quality prairie remnants around
Illinois (Figure 10.7). What type of leaf arrangement
Figure 10.6 Dehiscing follicles of Asclepias syriaca. do they have? Also, how would you describe the leaf
attachment for each specimen?

Asclepias verticillata L. (Whorled Milkweed)

Unlike many other species of milkweed, the leaves of


A. verticillata are very slender and display a whorled
arrangement; hence the species name ‘verticillata’ or
‘having whorls’.

Figure 10.7 Asclepias sullivantii plant with milky sap.

196 Integrative Biology 335: Plant Systematics


SOLANACEAE
potato family A5
♂ Ca⑤ Co⑤ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS

▪▪ Have PLICATE (folded) corolla


▪▪ Anthers often CONNIVENT

Solanum lycopersicum L. (Garden Tomato)

To observe the floral morphology characteristic of


Solanaceae, examine and dissect a S. lycopersicum
flower.

1. Examine a flower (Figure 10.8). What is its symmetry?

Figure 10.8 Close up of a Solanum lycopersicum, or


tomato, flower.

2. CALYX: How many sepals make up the calyx? Are


they distinct or connate?

3. COROLLA: How many petals make up the flower’s


PLICATE COROLLA (folded, connate petals)?

4. ANDROECIUM: Examine the epipetalous stamens with


their short filaments and elongate, tapering anthers
that terminate in a sterile tip. These CONNIVENT
anthers are just stuck together, not truly connate
(Figure 10.9). Although the anthers dehisce by
longitudinal slits, the pollen is released towards the
inside of the “tube”. How does it escape the flower?

Figure 10.9 Longitudinal section of a Solanum 5. GYNOECIUM: The ovaries of Solanaceae flowers are
lycopersicum flower. Notice the PLICATE
bicarpellate and often bilocular, although in many
corolla and the stamen’s CONNIVENT anthers.
cultivated varieties of tomato often more than 2
locules exist. Numerous ovules should be apparent
on axile placentae within the ovary.

6. DRAWING PORTFOLIO: Draw a floral diagram of a

Lab 10 - Asclepiadaceae, Solanaceae, Oleaceae and Caprifoliaceae 197


longitudinal section (L.S.) of S. lycopersicum, making
Box 1
sure to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

FRUIT TYPE

▪▪ Fruit is a berry or capsule

Capsicum annuum L. (Bell Pepper)


Solanum lycopersicum L. (Garden Tomato)
Physalis philadelphica Lam. (Tomatillo)
Solanum melongena L. (Eggplant)

Examine each of the edible berry fruits provided and


locate their persistent sepals. Do these berries develop
from a superior or an inferior ovary and how can you tell?
Sketch out a cross section of a tomato in Box 1 and label
the placentation type.

Solanum dulcamara L. (Climbing or Bittersweet


Nightshade)

This species of Solanum produces poisonous berry fruits,


though generally toxicity is highest in unripe berries
Figure 10.10 The poisonous fruits of Solanum (Figure 10.10). Incidences of poisoning in livestock,
dulcamara, or bittersweet nightshade.
however, are low due to the unpleasant odor that the plant
gives off.

Datura stramonium L. (Datura)

Datura plants are also extremely poisonous (Figure 10.11).


Their fruit is a 2-carpellate, 4-locular, spiny capsule that
opens by 4 apical valves. Based on this information, what
type of gynoecium does this fruit arise from?

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit of plants is mostly herbaceous


▪▪ Leaves have alternate arrangement

Solanum tuberosum L. (Potato)

Potatoes are actually a type of stem. What stem type are


Figure 10.11 A capsule fruit of Datura stramonium. they?

198 Integrative Biology 335: Plant Systematics


OLEACEAE
olive family A2
♂ Ca④ Co④ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS

▪▪ 4-merous flowers
▪▪ 2 stamens in Androecium
▪▪ 2-locular with two axile ovules in each locule (except 4-10
in Forsythia)

Forsythia spp. (Showy Forsythia)

To observe the floral morphology characteristic of


Oleaceae, you will examine a Forsythia flower.

1. Examine a flower (Figure 10.12). How would you


describe its floral symmetry?

2. CALYX: The calyx is deeply 4-lobed, made up of 4


Figure 10.12 A branch of Forsythia flowers.
connate sepals.

3. COROLLA: The corolla is made up of 4 connate petals.


What other term can be used to describe connate
petals?

4. ANDROECIUM: Observe the 2 EPIPETALOUS stamens


in the androecium (Figure 10.13). Few plant families
only have 2 stamens!

5. GYNOECIUM: The ovary is superior with a terminal


style and a 2-lobed stigma. Thus, two connate carpels
make up the flower’s syncarpous gynoecium. In
Forsythia, there are 4-10 ovules per locule.

6. DRAWING PORTFOLIO: Draw a floral diagram of a


longitudinal section (L.S.) of Forsythia, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

Figure 10.13 Longitudinal section of a Forsythia flower.

Lab 10 - Asclepiadaceae, Solanaceae, Oleaceae and Caprifoliaceae 199


Fraxinus pennsylvanica Marshall (Green Ash)

In the Oleaceae family there is a trend towards wind


pollinated flowers. For those plants with imperfect flowers,
like Fraxinus, the plants are dioecious (Figure 10.14). The
flowers of ash are very reduced. Being wind pollinated,
there is no need for a perianth to attract pollinators, so
the calyx and corolla are absent. Be sure to examine the
herbarium specimens of the staminate and carpellate trees
on display.

Figure 10.14 Fraxinus pennsylvanica flowers.


FRUIT TYPE

▪▪ Fruits variable - can be a berry, capsule, drupe or samara

Ligustrum vulgare L. (Wild Privet or Common Privet)

Examine the tiny berries produced by L. vulgare. Seed


dispersal occurs via thrushes, who eat the berries
and release the seeds in new locations through their
droppings. Don’t be fooled into eating the berries yourself
though, they are toxic to humans!

Syringa vulgaris L. (Common Lilac)

This species produces capsule fruits, which dehisce


to release two winged seeds. What type of capsule is
it classified as? Also, what is the inflorescence type
Figure 10.15 Drupe fruit of Olea europea. assuming that each capsule fruit was originally a flower?

Olea europaea L. (Olive)

Olives are botanically considered drupes (Figure 10.15).


After looking at the dissected olives on display, explain
why they are classified as drupes and not berries.

Fraxinus pennsylvanica Marshall (Green Ash)

Ash trees produce an overwhelming amount of samaras


(Figure 10.16). How are these fruits dispersed?

Figure 10.16 Samaras produced by a Fraxinus tree.

200 Integrative Biology 335: Plant Systematics


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit is either a tree or shrub


▪▪ Leaves have opposite arrangement

Fraxinus pennsylvanica Marshall (Green Ash)


Syringa vulgaris L. (Common Lilac)

Using the “Dichotomous Key for Woody Plants” in Lab 1:


Woody Twig Terminology, determine which twig is from F.
pennsylvanica and which is from S. vulgaris. Notice how
the leaf scars on the twigs appear opposite one another
due to the opposite leaf arrangement characteristic of the
Oleaceae family.

Figure 10.17 Leaves of a Fraxinus quadrangulata, or


blue ash tree.

Lab 10 - Asclepiadaceae, Solanaceae, Oleaceae and Caprifoliaceae 201


CAPRIFOLIACEAE
A 5
honeysuckle family ♂ Ca 5 Co⑤

Eudicots: Asterid Clade G 2-5

FLORAL CHARACTERISTICS

▪▪ Ovaries of two flowers often paired


▪▪ Ovary inferior with 2-5 locules

Kolkwitzia amabilis Graebn. (Beautybush)


Lonicera japonica Thunb. (Japanese Honeysuckle)
Lonicera sempervirens L. (Trumpet Honeysuckle)
Lonicera tatarica L. (Tatarian Honeysuckle)
Viburnum carlesii Hemsl. (ADOXACEAE) (Korean Spice)
Viburnum opulus L. (ADOXACEAE) (European
Cranberrybush)
Figure 10.18 Flowers of Lonicera japonica. Flowers in the Caprifoliaceae family have inferior ovaries,
and often the ovaries of two separate flowers will be paired
together. To see these features, examine the herbarium
specimens and the dissected flower diagrams provided.
What term is used to describe the stamens in the family
(as well as most of the Asterid clade)?

These showy flowers are pollinated by various nectar-


gathering insects (primarily bees and wasps) and birds.
What is the floral symmetry of the flowers in Figure 10.18?
What about Figure 10.19?

FRUIT TYPE

Figure 10.19 Flowers of Viburnum opulus.


HABIT AND VEGETATIVE CHARACTERISTICS

202 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruits can be berries, capsules or drupes

Lonicera maackii Maxim. (Amur Honeysuckle)


Viburnum spp. (ADOXACEAE)

All of the members of the genus Lonicera produce berries,


which will often be paired like the ovaries they arose from
(Figure 10.20). The color of these berries can be red, blue
or black and each fruit will contain a number of seeds.

Alternatively, the members of the genus Viburnum produce


single-seeded drupe fruits (Figure 10.21,) although the
color varies (i.e. red, purple, blue or black). Check out the
herbarium specimens to see the drupes produced by a few
different Viburnum species. What is the major difference
between a berry and a drupe?

Figure 10.20 Fruit of Lonicera purpusii.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Plants possess a variety of woody habit types


▪▪ Leaves have opposite arrangement and do not have
stipules

Figure 10.21 Fruit of Viburnum opulus.


Lonicera maackii Maxim. (Amur Honeysuckle)

Review your vegetative terminology while observing this


herbarium specimen.

Complexity __________________________________

Arrangement __________________________________

Attachment __________________________________

Venation __________________________________

Blade shape __________________________________

Leaf apex __________________________________

Leaf base __________________________________


Figure 10.22 Close up of Lonicera maackii flowers. Leaf margin __________________________________

Lab 10 - Asclepiadaceae, Solanaceae, Oleaceae and Caprifoliaceae 203


204 Integrative Biology 335: Plant Systematics
LABORATORY 11
SCROPHULARIACEAE, LAMIACEAE AND ASTERACEAE

PHYLOGENETIC RELATIONSHIPS

The order Lamiales consists of about 24 families


and 17,800 species and is classified within the
Core Asterid clade (Lamiids subclade). Included
here are Scrophulariaceae and Lamiaceae,
which we will study in today’s lab. The order is
undoubtedly monophyletic, being held together
by the shared possession of a large suite
of morphological, anatomical, chemical and
molecular characters.

Scrophulariaceae, as traditionally delimited,


is highly polyphyletic, with some members of
this group more closely related to members of
various other families of Lamiales than they are
to other scrophs. Recent systems of classification
place the following genera formerly found in the
Scrophulariaceae family in the Plantaginaceae
family: Antirrhinum, Digitalis, Mimulus,
Penstemon, Plantago and Veronica. The family
Scrophulariaceae is treated in a very strict sense
and only includes a few genera (i.e. Verbascum
and Scrophularia, which occur in North America).
In this class, we will treat the family in its traditional (i.e., broad) sense to include those genera recently placed in the
Plantaginaceae. Scrophulariaceae is well known for its many ornamental plants.

The Asteraceae family is contained within the order Asterales, which includes 12 families and about 25,000 species.
Asterales is found in the Campanulids subclade.

Lab 11 - Scrophulariaceae, Lamiaceae and Asteraceae 205


SCROPHULARIACEAE
figwort family A2 or 4 (5)
♂ CaZ⑤ CoZ⑤ G②

Eudicots: Asterid Clade

FLORAL CHARACTERISTICS

▪▪ Zygomorphic symmetry due to 2-lipped to nearly regular


corolla
▪▪ DIDYNAMOUS stamens (2 or 2+2) sometimes with fifth
staminode (STAMINODIUM) present

Antirrhinum majus L. (Snapdragons)

To observe the floral morphology characteristic of


Scrophulariaceae, you will examine and dissect a A. majus
flower. If you hold a fresh blossom by the corolla tube
immediately below the lobes and squeeze gently between
thumb and forefinger, the two lips will open like the jaws
of an animal’s mouth (a dragon perhaps?) and will snap
closed again when the pressure is released. Hence the
common name for the species, ‘snapdragon’.

1. Observe a flower (Figure 11.1). What is its symmetry?

2. CALYX: The calyx of these flowers is deeply 5-parted


and zygomorphic.
Figure 11.1 Antirrhinum majus inflorescence.
3. COROLLA: The corolla is sympetalous and BILABIATE
(upper lip is formed from 2 petals, while the lower lip
is formed from 3 petals). Observe that the lower lip
has a prominent palate that closes the throat of the
corolla.

4. ANDROECIUM: How many stamens are there? In


Scrophulariaceae, the stamens are arranged in a
DIDYNAMOUS fashion (Figure 11.2). What other plant
family covered this semester had TETRADYNAMOUS
stamens? What is the difference between these two
terms?

5. GYNOECIUM: Examine the syncarpous gynoecium of


this flower consisting of 2 connate carpels. Its superior
Figure 11.2 Longitudinal section of an Antirrhinum ovary is bilocular and contains numerous axile ovules.
majus flower. The style is terminal, as it is in all the other families
we have studied so far.

206 Integrative Biology 335: Plant Systematics


6. DRAWING PORTFOLIO: Draw a floral diagram of a
longitudinal section (L.S.) of A. majus, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.

Penstemon spp. (Beardstongue)

Flowers from this genus possess a BILABIATE corolla,


DIDYNAMOUS stamens (2 + 2), a gynoecium with a 2-lobed
stigma and a superior ovary (Figure 11.3). They also have
Figure 11.3 Longitudinal section of a Penstemon
pallidus flower. a 5th hairy, sterile stamen (hence the common name
‘beardstongue’). What is the name of this structure?

FRUIT TYPE

▪▪ Fruit is a 2-valved capsule

Penstemon spp. (Beardstongue)


Verbascum blattaria L. (Moth Mullein)
Verbascum thapsus L. (Common Mullein)

Observe the Scrophulariaceae fruits provided. These


capsule fruits have two valves which arise from the two
Capsules produced by a Penstemon plant.
connate carpels that make up the syncarpous gynoecium
Figure 11.4
of the original flower (Figure 11.4).

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Many photosynthetic root parasites

Pedicularis canadensis L. (Canadian Lousewort)

This species is a hemiparasite, meaning that it establishes


connections with other plants’ roots to obtain resources
but it is also capable of photosynthesis itself. Its habit is
either an herb or a subshrub. The fruits of P. canadensis
are capsules that only open on the upper side; they are an
example of “splash-cup” dispersal (Figure 11.5).

Figure 11.5 Capsules produced by Pedicularis


canadensis.

Lab 11 - Scrophulariaceae, Lamiaceae and Asteraceae 207


LAMIACEAE
mint family A4
♂ CaZ⑤ CoZ⑤ G②

Eudicots: Asterid Clade

INFLORESCENCE TYPE

▪▪ Axillary cymes or VERTICILS

Mentha spicata L. (Spearmint)


Nepeta cataria L. (Catnip)
Stachys arenicola Britt. (Hairy Hedgenettle)

Observe how the flowers are arranged in VERTICILS or


whorls in these plant species (Figure 11.6). This is a
diagnostic inflorescence feature of many members of
Lamiaceae.

Figure 11.6 Teucrium canadense flowers arranged in


VERTICILS.

FLORAL CHARACTERISTICS

▪▪ Zygomorphic symmetry due to BILABIATE corolla (upper


portion of corolla has 2-lobes and lower part has 3-lobes)
▪▪ DIDYNAMOUS stamens (2+2 arrangement)
▪▪ Gynoecium deeply 4-lobed with GYNOBASIC style and 1
basal/axile ovule per locule

Lavendula spp. (Lavendar)

To observe the floral morphology characteristic of


Lamiaceae, you will examine a Lavendula flower.

1. Examine a flower. What is its symmetry?

2. CALYX: How many sepals make up the calyx and are


they distinct or connate?

Figure 11.7 Base of a Physostegia virginiana flower.

3. COROLLA: The corolla is made up of five connate


petals and is described as BILABIATE since it has two
lips, a 2-lobed upper lip and a 3-lobed lower lip.

208 Integrative Biology 335: Plant Systematics


4. ANDROECIUM: Observe the EPIPETALOUS stamens
with their DIDYNAMOUS arrangement. How many
stamens make up the androecium?

5. GYNOECIUM: How many carpels make up the


syncarpous gynoecium of this flower? Examine
the ovary of the flower. It is deeply four lobed and
possesses a GYNOBASIC style (Figure 11.7 and 11.8).
What does GYNOBASIC mean?

Molucella laevis L. (Bells-of-Ireland)


Salvia spp. (Sage)
Teucrium spp. (Germander)

Dissect one of the Molucella flowers and check out the


Figure 11.8 Dissected parts of a Teucrium canadense photos and herbarium specimens provided for Salvia
flower including its 4-lobed ovary. and Teucrium. As you do so, try to pick out the important
Lamiaceae floral characteristics.BIT AND VEGETATIVE
CHARACTERISTICS

FRUIT TYPE

▪▪ Produces 4 nutlets

Leonurus cardiaca L. (Common Motherwort)

Take one of the fruits and dissect it so that you can see
Figure 11.9 Nutlets of Leonurus cardiaca. the 4 nutlets that each flower produces (Figure 11.9). What
is the difference between a nutlet and a nut?

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Stems are square in cross section


▪▪ Leaves are simple and oppositely arranged
▪▪ Produce essential oils and are aromatic

Majorana spp. (Majoram)


Mentha spp. (Mint)
Figure 11.10 Bee visiting a Monarda fistulosa flower. Ocimum spp. (Basil)

Lab 11 - Scrophulariaceae, Lamiaceae and Asteraceae 209


Thymus spp. (Thyme)
Box 1
Take some time to observe the vegetative characteristics
of these plants. What is their habit?

Cut a small piece of stem off of one of the plants and


draw its cross section in Box 1. The cross section shape
is a very important feature for the Lamiaceae family! Also
notice the plants’ lovely aromatic scents.

210 Integrative Biology 335: Plant Systematics


ASTERACEAE
A 5
sunflower family ♂ CaP Co⑤

Eudicots: Asterid Clade G②

INFLORESCENCE TYPE

▪▪ Head (CAPITULUM, CAPITULA) inflorescences surrounded


by PHYLLARIES
▪▪ Three CAPITULUM types: RADIATE CAPITULUM, DISCOID
CAPITULUM and LIGULATE CAPITULUM

When looking at a sunflower, blazing star or dandelion,


you might assume you are looking at a solitary flower
(Figures 11.11 and 11.12). However, that is not the case.
If you were to split one of the “flowers” lengthwise, you
will see many small flowers (called FLORETS) arranged
side by side on a large common receptacle. (Note: The
use of the word “receptacle” is technically not correct, as
a true receptacle is part of a flower, while in this instance
it is used for the base of an inflorescence.) This compact
arrangement of sessile flowers is a called a CAPITULUM
(HEAD) or composite head, hence the alternative name
for Asteraceae, Compositae. Asteraceae CAPITULA are
Figure 11.11 Field of Helianthus annuus flowers. divided into three types based on the type of FLORETS that
make them up. The CAPITULA are surrounded by several
to many green bracts called PHYLLARIES or involucral
bracts. The stalk of the inflorescence (i.e. CAPITULUM) is
called a PEDUNCLE. Is a capitulum a determinate or an
indeterminate inflorescence?

FLORAL CHARACTERISTICS

▪▪ Two kinds of flowers (FLORETS): DISK and RAY


▪▪ Bracts called CHAFF and calyx called a PAPPUS
▪▪ Stamens with distinct filaments attached to base of
corolla, but anthers connate into a tube around style
▪▪ Syncarpous gynoecium is made up of 2 connate carpels
and has a 1-locular, inferior ovary with 1 basal ovule

There are two kinds of flowers (FLORETS):

1. DISK: actinomorphic symmetry with a tubular corolla


Figure 11.12 Radiate capitulum of Helianthus annuus. 2. RAY: zygomorphic symmetry with a ligulate corolla

Lab 11 - Scrophulariaceae, Lamiaceae and Asteraceae 211


As previously mentioned, the three CAPITULUM types are
differentiated based on which FLORET(S) they possess.

1. RADIATE CAPITULUM: both DISK and RAY FLORETS


2. DISCOID CAPITULUM: only DISK FLORETS
3. LIGULATE CAPITULUM: only RAY FLORETS

RADIATE CAPITULUM
Helianthus spp. (Sunflower)

The genus name Helianthus comes from the Greek words


Figure 11.13 Radiate capitulum of Helianthus
angustifolius with disk and ray florets. “helios” (sun) and “anthos” (flower). Pretty isn’t it? Now
examine the RADIATE CAPITULUM of Helianthus, with its
inner DISK FLORETS (actinomorphic and brown) and outer
RAY FLORETS (zygomorphic, strap-like and yellow; Figures
11.13 and 11.14).

DISK FLORETS
1. Identify the CHAFF, if present. These are the bracts
that subtend the florets and arise off the receptacle.

2. CALYX: Find the PAPPUS (the modified calyx). The


pappus may be modified into long white bristles,
retrorsely barbed awns, scales, or be missing
altogether.

3. COROLLA: Observe that the corolla is actinomorphic,


Figure 11.14 Longitudinal section of a Helianthus tubular and 5-lobed at the summit.
angustifolius radiate capitulum.
4. ANDROECIUM: The androecium is made up of 5
stamens. The stamens’ anthers are united by their
margins to form a cylinder around the style, while the
filaments are distinct but adnate to the corolla tube
(Figure 11.15). Based on this, what term can be used
to describe the stamens?

5. GYNOECIUM: Observe the 2 stigmas, elongate style


and inferior ovary of the syncarpous gynoecium. Each
ovary contains only one locule and one basal ovule.
What is the insertion type?

RAY FLORETS
1. Identify the CHAFF if present.
Figure 11.15 Close up of disk and ray florets from
Heliopsis helianthoides. 2. CALYX: Find the PAPPUS (the modified calyx).

3. COROLLA: Observe that the zygomorphic corolla


arises as a very short tube at the summit of the ovary
but immediately splits into a long, narrow, flat, strap-

212 Integrative Biology 335: Plant Systematics


shaped structure with commonly three terminal lobes.

4. ANDROECIUM: In this floret there is no androecium


(therefore, the florets are imperfect) or, in some
species, the florets may be sterile. If sterile, what is
their function?

5. GYNOECIUM: Observe the inferior ovary of the


gynoecium.

6. DRAWING PORTFOLIO: Draw a floral diagram of a


longitudinal section (L.S.) of a Helianthus CAPITULUM,
making sure to include its floral formula. On the same
page, include floral diagrams of individual DISK and
RAY FLORETS. Follow the format as instructed in Lab 3:
Figure 11.16 Liatris pycnostachya discoid capitulum. Floral Terminology.

Chrysanthemum spp. (Daisy)

Take time to dissect a Chrysanthemum CAPITULUM and


try to pick out the same features that you saw in the
Helianthus CAPITULUM including the two types of FLORETS
and their constituent parts.

DISCOID CAPITULUM
Liatris spp. (Blazing Star)

DISCOID CAPITULA are inflorescences that contain only DISK


FLORETS (Figure 11.16). Examine one to verify that their
structure is the same as DISK FLORETS found in a RADIATE
Figure 11.17 Liatris pycnostachya disk floret. CAPITULUM (Figure 11.17).

LIGULATE CAPITULUM
Taraxacum officinale F.H. Wigg (Common Dandelion)

LIGULATE CAPITULA are inflorescences that contain only


RAY FLORETS (Figure 11.18). These RAY FLORETS are
somewhat different than those found on the margin of a
RADIATE CAPITULUM. Most notably they are perfect, unlike
RAY FLORETS in a RADIATE CAPITULUM which are either
carpellate only or sterile. Dissect a T. officinale flower to
see the RAY FLORET features.

Figure 11.18 Ligulate head of Taraxacum officinale. 1. Observe a RAY FLORET. What is its floral symmetry?

Lab 11 - Scrophulariaceae, Lamiaceae and Asteraceae 213


2. COROLLA: The corollas of the outer florets are
obviously flat. Those of the center appear regular and
might be considered tubular. However they are the
same as the outer ones, only they are younger and
have not opened or expanded. There are 5 terminal
lobes to the floret corolla (Figure 11.19).

3. ANDROECIUM: In this floret the filaments are distinct


above the point where they attach to the corolla tube,
but the anthers are connate into a tube.

4. GYNOECIUM: Observe the inferior ovary, solitary style


and 2 stigma lobes of the gynoecium (Figure 11.20).

POLLINATION MECHANISM

Figure 11.19 Taraxacum officinale ray floret from a


ligulate head. In the Asteraceae family, pollen is shed inside the staminal
tube. As the style of the gynoecium elongates, the pollen is
pushed out the tip of the staminal tube. Sometimes there
is a special “brush” towards the tip of the style to assist
in pushing the pollen out. The 2 stigmas are folded shut
as the style elongates, but they spread open after they
emerge beyond the anther tube. Now pollination is free to
occur.

FRUIT TYPE

▪▪ Fruit is an achene

RADIATE CAPITULUM: Helianthus spp. (Sunflower)

The fruit type typically produced by Asteraceae is an


Figure 11.20 Details of Taraxacum officinale ray florets. achene (Figure 11.21). What are some key features of
an achene fruit based on the examples provided (i.e.
characteristics of parent flower, fruit features, etc.)?

Figure 11.21 Achenes from Helianthus disk florets.

214 Integrative Biology 335: Plant Systematics


DISCOID CAPITULUM: Liatris spp. (Blazing Star)
LIGULATE CAPITULUM: Tragopogon spp. (Goat’s Beard)

CYPSELA fruits are restricted to some species in


Asteraceae. They are dry, indehiscent and very similar to
ACHENES except that they usually have an adnate calyx.
In Liatris, the calyx is modified into bristle-like appendages
called the PAPPUS (Figure 11.22).

Note: A sunflower fruit (commonly referred to as a


sunflower seed) is correctly termed a CYPSELA but does
not possess an adnate calyx. Consequently, it can also be
referred to as an ACHENE.

Figure 11.22 Achene from Liatris aspera.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit is usually an herb

RADIATE CAPITULUM: Tagetes spp. (Marigold)

Like this Tagetes species, the majority of the Asteraceae


family members are herbaceous. If they are available,
dissect the RADIATE CAPITULA of the Tagetes species.

LIGULATE CAPITULUM: Taraxacum officinale F.H. Wigg


(Common Dandelion)

Many members of the Asteraceae with a ligulate capitulum


Figure 11.23 Taraxacum officinale plant. have milky sap, like the T. officianale on display (Figure
11.23). What root type does this plant exhibit?

Lab 11 - Scrophulariaceae, Lamiaceae and Asteraceae 215


216 Integrative Biology 335: Plant Systematics
LABORATORY 12
ROSACEAE, ACERACEAE AND APIACEAE

PHYLOGENETIC RELATIONSHIPS

In today’s Lab we will examine Rosaceae and


Aceraceae from the Rosid clade, and Apiaceae
from the Asterid clade.

Rosaceae is treated in the Rosales clade of the


Rosids. Many characters support its monophyly,
such as the presence of numerous stamens,
the absence of alkaloids and DNA sequences.
Alternate leaves with stipules plus showy flowers
with radial symmetry and a hypanthium also aid
in their recognition. Traditionally, the family is
divided into four subfamilies, and we will highlight
these subfamilies in Lab. Fruit type, carpel
number, ovary position and base chromosome
number are the primary criteria in subfamily
recognition. Recent results of phylogenetic
analyses of molecular data, however, suggest
that not all of these subfamilies are monophyletic.

Aceraceae is now treated within the family


Sapindaceae (the soapberry family). Along
with the maples and relatives are members of
the Hippocastanaceae (horse chestnuts and
relatives) and several important tropical fruits,
such as longan, lychee and rambutan (in which
the fleshy part of the fruit you eat is derived
from a large aril surrounding the seed). While
your textbook defines the Sapindaceae broadly,
we will treat the Aceraceae as a separate,
distinct family (as a matter of convenience
only). Therefore, the Aceracee sensu stricto
(Aceraceae s.s.) is circumscribed by plants
bearing opposite leaves, non-appendaged petals,
stamens borne on a nectar disk, and two ovules
per carpel. Aceraceae s.s. is classified within the
order Sapindales of the Malvids subclade. So,
actually this family is more closely related to the
Malvaceae and Brassicaceae than it is to the
Apiaceae.

The family Apiaceae (or Umbelliferae, the carrot


family), as circumscribed in the Judd et al. text,
is treated in the broad sense to include not only
those herbaceous species traditionally considered in

Lab 12 - Rosaceae, Aceraceae and Apiaceae 217


Apiaceae, but also those woody species which are usually segregated as a separate family, the Araliaceae. However,
in this class, we consider only Apiaceae sensu stricto (Apiaceae s.s.) as constituting the family. Synapomorphies for
Apiaceae s.s. include hollow internodes, secretary canals containing ethereal oils and resins, umbel inflorescences,
a stylopodium atop the inferior ovary, and a schizocarpic fruit. The family Apiaceae s.s is treated within the order
Apiales of the Campanulids subclade (of the core Asterid clade), as indicated by their ovules with a single integument
and usually a thin megasporangium wall, sympetalous corollas, 5 stamens and molecular data. This order is closely
related to the Asterales (e.g. Asteraceae) and Dipsacales (e.g. Caprifoliaceae). Apiaceae s.s. is the subject of Dr.
Downie’s research.

218 Integrative Biology 335: Plant Systematics


ROSACEAE
rose family

♀ Ca5 Co5 A∞ G ∞ or G1 or G 2-5
Eudicots: Rosid Clade

FOUR SUBFAMILIES

▪▪ SPIRAEOIDEAE
▪▪ ROSOIDEAE
▪▪ AMYGDALOIDEAE
▪▪ MALOIDEAE

Rosaceae is divided into four distinct subfamilies.


The subfamilies can be differentiated based on carpel
number and fusion, ovary position, fruit type and base
chromosome number.

Figure 12.1 Bowl of Rosaceae fruit. As Dr. Ken


Robertson says, “The Rose Family FLORAL CHARACTERISTICS
(Rosaceae) makes life worth living!”
▪▪ Actinomorphic symmetry
▪▪ Flowers with HYPANTHIUM

Though all members of the Rosaceae family have perfect,


actinomorphic flowers with a hypanthium, the subfamilies
differ in their carpel number and fusion as well as their
ovary position.

Subfamily Floral Formula Carpel Number and Fusion Ovary Position

SPIRAEOIDEAE ♂
♀ Ca5 Co5 A10-∞ G 2-5 Few (5) distinct carpels Superior ovary
(Spirea Subfamily)

ROSOIDEAE ♂
♀ Ca5 Co5 A10-∞ G ∞ Many distinct carpels Superior ovary
(Rose Subfamily)

AMYGDALOIDEAE ♂
♀ Ca5 Co5 A5-15 G1 1 carpel Superior ovary
(Peach Subfamily)


♀ Ca5 Co5 A10-∞
MALOIDEAE 2-5 carpels that are fused to
Inferior ovary
(Apple Subfamily) hypanthium
G 2-5

Lab 12 - Rosaceae, Aceraceae and Apiaceae 219


SPIRAEOIDEAE
Spiraea spp. (Spirea)
Exochorda racemosa (Lindl.) Rehder (Pearlbush)

To examine the floral characteristics of Spiraeoideae,


dissect either a Spiraea (Figure 12.2) or an Exochorda
racemosa flower (Figure 12.3).

1. Observe a flower. What is its symmetry?


Figure 12.2 Cluster of Spirea flowers.

2. CALYX, COROLLA and ANDROECIUM: These three floral


series are adnate to form what?

3. GYNOECIUM: Five weakly united carpels make up


the gynoecium of this flower, making it atypical
of subfamily Spiraeoideae (Figure 12.4). Most
Spiraeoideae species have apocarpous gynoecia
made up of 2-5 distinct carpels. What is the ovary
Figure 12.3 Flower of Exochorda racemosa.
position?

4. In Box 1, sketch a longitudinal section (L.S.) of a


Spiraea or an Exochorda racemosa flower.

5. Construct the floral formula for the flower you are


dissecting in the space below.

Figure 12.4 Longitudinal section of an Exochorda


racemosa flower.

Box 1

220 Integrative Biology 335: Plant Systematics


ROSOIDEAE
Duchesnea spp. (Duchesnea)
Fragaria spp. (Strawberry)
Rosa spp. (Rose)

Subfamily Rosoideae species are distinguished from


members of the other three subfamilies by their numerous
and distinct carpels and superior ovaries (Figure 12.5).
Dissect one of the species on display.

1. What is the floral symmetry of the flower?

Figure 12.5 Flower of Rosa carolina.

2. CALYX, COROLLA and ANDROECIUM: How many


sepals, petals and stamens does your flower
possess? In Rosa, a deep hypanthium is present, and
though it may appear to be, the hypanthium is NOT
adnate to the gynoecium (Figure 12.6).

3. GYNOECIUM: Observe how the carpel styles protrude


through the opening in the top of the hypanthium.
What is the gynoecium type if it is made up of many
distinct carpels? Also, what is the ovary position and
insertion type in the flower?

4. In Box 2, sketch a longitudinal section (L.S.) of a


Duchesnea, Fragaria or Rosa flower.

5. Construct the floral formula for the flower you are


dissecting in the space below.
Figure 12.6 Longitudinal section of a Rosa rubrifolia
flower.

Box 2

Lab 12 - Rosaceae, Aceraceae and Apiaceae 221


AMYGDALOIDEAE
Prunus spp. (Plum)

To learn the floral morphology characteristic of Rosaceae


subfamily Amygdaloideae, you will examine and dissect
Prunus flowers.

1. Observe a flower (Figure 12.7). What is its floral


symmetry?

2. CALYX, COROLLA and ANDROECIUM: Prunus flowers


have large hypanthiums lined with nectar secreting
tissue that are formed from the adnation of these
three floral series (Figure 12.8). How may sepals,
petals and stamens does your flower possess?

Figure 12.7 Inflorescence of Prunus padus.


3. GYNOECIUM: At the base of the hypanthium is a
monocarpous gynoecium with a superior ovary. What
is the insertion type?

4. In Box 3, sketch a longitudinal section (L.S.) of a


Prunus flower.

5. Construct the floral formula for the Prunus flower you


are dissecting in the space below.

Figure 12.8 Longitudinal section of a Prunus padus


flower.

Box 3

222 Integrative Biology 335: Plant Systematics


MALOIDEAE
Pyrus calleryana ‘Bradford’ (Pear)
Malus spp. (Apple)

To learn the floral morphology characteristic of Subfamily


Maloideae, you will examine and dissect Pyrus and Malus
flowers. The genus names Pyrus and Malus come from
the Latin for “pear” and “apple,” respectively.

1. Observe a flower (Figure 12.9). What is its floral


symmetry?

Figure 12.9 Cluster of Malus flowers.

2. CALYX, COROLLA and ANDROECIUM: Like the other


Rosaceae subfamilies, Maloideae flowers have a
hypanthium (Figure 12.10). How many sepals, petals
and stamens do they possess?

3. GYNOECIUM: What is the gynoecium type, ovary


position and insertion type of these flowers?

4. In Box 4, sketch a longitudinal section (L.S.) of a


Pyrus calleryana or Malus flower.

5. Construct the floral formula for the flower you are


dissecting in the space below.

Figure 12.10 Longitudinal section of a Malus flower.

Box 4

Lab 12 - Rosaceae, Aceraceae and Apiaceae 223


FRUIT TYPE

▪▪ Do NOT produce capsules or berries

The fruit types produced in the Rosaceae family vary


among the four subfamilies. In fact, fruit type is one of the
major features that distinguish the subfamilies from one
another.

Subfamily Fruit Type


Figure 12.11 Spirea × vanhouttei flowers produce an
aggregate fruit of follicles. SPIRAEOIDEAE Follicle

ROSOIDEAE Achenes (in a HIP) or drupelets

AMYGDALOIDEAE Drupe

MALOIDEAE POME

SPIRAEOIDEAE
Exochorda giraldii Hesse (Redbud Pearlbush)
Figure 12.12 Arrow is pointing to an achene fruit of Physocarpus opulifolius (L.) Maxim. (Ninebark)
Fragaria × ananassa. Spiraea spp. (Meadowsweet)

Follicles, a dry dehiscent fruit type, are characteristic of the


Spiraeoideae subfamily (Figure 12.11). Examine the array
of follicle fruits on display. What gynoecium type must
Spiraeoideae species have in order to produce them?

ROSOIDEAE
Fragaria × ananassa (Strawberry Hybrid)

The Rosoideae subfamily typically produces achene fruits.


In Fragaria × ananassa, the fruit is composed of many
achenes on the outside of an enlarged receptacle (Figure
12.12). Thus, what three fruit types can a strawberry be
classified as?

Rosa canina L. (Dog Rose)


Rosa multiflora Thunb. (Multiflora Rose)

Rosa species produce HIPS containing many achene fruits


(Figure 12.13). The HIP itself is formed when the original
flower’s deep hypanthium becomes fleshy and its ovaries
Figure 12.13 Hip of Rosa setigera. (located inside the hypanthium) mature into achenes. Cut

224 Integrative Biology 335: Plant Systematics


open one of the HIPS provided and examine the achenes.

Rubus idaeus L. (American Red Raspberry)


Rubus allegheniensis Porter. (Allegheny Raspberry)

Although less common, the Rosoideae subfamily does


also produce drupelet fruits. For example, the genus
Rubus produces “AGGREGATE FRUITS OF DRUPELETS”
(Figure 12.14). What type of gynoecium is required to
produce this fruit type?

Figure 12.14 Rubus aggregate fruit of drupelets.

AMYGDALOIDEAE
Prunus americana Marsh (American Plum)
Prunus armeniaca L. (Apricot)
Prunus avium (L.) L. (Sweet Cherry)
Prunus dulcis (Mill.) D.A. Webb (Sweet Almond)
Prunus persica (L.) Batsch (Peach/Nectarine)

After pollination, the petals of Amygdaloideae flowers fall


off quickly, while the sepals and stamens persist for a short
while. The carpel then enlarges and begins to develop into
Figure 12.15 Drupe fruit of Prunus padus. a fruit (Figure 12.15). What are some of the characteristics
that define the drupe fruit type?

MALOIDEAE
Amelanchier spp. (Serviceberry)
Crataegus mollis Scheele (Downy Hawthorn)
Malus pumila Mill. (Paradise Apple)
Pyrus communis L. (Common Pear)

A POME is the fleshy ACCESSORY FRUIT of subfamily


Figure 12.16 POME fruit of Malus pumila.
Maloideae and is derived from the adnation of the
hypanthium to the ovary wall. Thus, the flesh of apples
and pears is mostly hypanthium tissue (Figure 12.16). Did
the POMES on display arise from a superior or an inferior
ovary? How can you tell based on the fruits?

Figure 12.17 POME fruit of Malus ‘Red Jade’.

Lab 12 - Rosaceae, Aceraceae and Apiaceae 225


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Habit varies with Rosaceae subfamily

Look back at the material provided under the flower and


fruit portion of this lab to see the habits characteristic of
the four Rosaceae subfamilies.

Base
Subfamily Plant Habit
Chromosome #

Mostly shrubs, some


SPIRAEOIDEAE herbs
9

Shrubs and perennial


ROSOIDEAE herbs
7

AMYGDALOIDEAE Trees and shrubs 8


Figure 12.18 Trunk (with horizontal lenticels) of a Prunus
maackii tree. MALOIDEAE Trees and shrubs 17

226 Integrative Biology 335: Plant Systematics


ACERACEAE ♂
maple family ♂
♀ or ♀ Ca5(4) Co5(4) A8,10,5 * G② †
Eudicots: Rosid Clade Rudimentary in ♀ flowers (*) and ♂ flowers (†)

FLORAL CHARACTERISTICS

▪▪ Perfect or imperfect flowers that are wind or insect


pollinated
▪▪ Syncarpous gynoecium composed of two connate carpels
▪▪ Superior ovary is winged and 2-locular with 2 axile ovules
per locule

Acer platanoides L. (Norway Maple)

To observe the floral morphology characteristic of


Aceraceae, you will examine carpellate and staminate
A. platanoides flowers. This species is insect pollinated,
so you will notice that all flowers, regardless of gender,
Figure 12.19 Cluster of Acer platanoides flowers. possess a large, green nectar disk. Also, carpellate and
staminate flowers are typically located on the same tree
in A. platanoides (Figure 12.19). What then is the plant
condition?

CARPELLATE FLOWER
1. Examine a carpellate flower. What is its symmetry?

Figure 12.20 Acer platanoides female flower. 2. CALYX and COROLLA: How many sepals and petals
make up the calyx and corolla, respectively?

3. ANDROECIUM: Examine the staminodes present in the


female flower (Figure 12.20). How many are there and
what is their function?

4. GYNOECIUM: The syncarpous gynoecium of this flower


is made up of how many connate carpels? Note that
the ovary already shows signs of wing development;
Figure 12.21 Longitudinal section of an Acer platanoides at this stage of floral maturity, there is a small wing on
female flower. each side of the ovary (Figure 12.21).

Lab 12 - Rosaceae, Aceraceae and Apiaceae 227


STAMINATE FLOWER
1. Examine a staminate flower. What is its symmetry?

2. CALYX and COROLLA: How many sepals and petals


make up the calyx and corolla, respectively?

Figure 12.22 Acer platanoides male flower.


3. ANDROECIUM: Note that there are 8 stamens in the
androecium of this flower. Amongst plant families in
general this is an unusual number of stamens to have,
but it is common in the Aceraceae.

4. GYNOECIUM: There is no rudimentary gynoecium in


staminate flowers (Figure 12.22).

5. DRAWING PORTFOLIO: Draw a floral diagram of a


longitudinal section (L.S.) of the carpellate and
staminate A. platanoides flowers, making sure to
include their floral formulas. Follow the format as
instructed in Lab 3: Floral Terminology.

Figure 12.23 Male and female Acer saccharinum


inflorescences. Acer rubrum L. (Red Maple)

The flowers of maples range from comparatively showy


and insect-pollinated like A. platanoides to small,
inconspicuous and wind-pollinated. Many are intermediate
between the two. Examine the Acer rubrum flowers
provided, comparing their structure to what you observed
for A. platanoides. In the male flower there are 8 stamens
arising off a nectariferous disk. How can you tell that
these stamens are functional? - Check if they contain
pollen! These stamens surround a small, non-functional
gynoecium. In the female flower there is a large, glabrous,
bilocular ovary with 2 distinct styles. The ovary is visibly
compressed and has 2 projecting wing-like protuberances.
You can also see staminodes arising off a nectariferous
disk. Based on this description and your own observations,
what is their pollination mechanism?

Acer saccharinum L. (Silver Maple)

The flowers of A. saccharinum are adapted to wind


pollination rather than insect pollination. This species
blooms very early in the spring before the leaves appear.
The female flowers have 2 long styles that project outward
to catch windblown pollen. Note the staminodes in the
female flowers. The male flowers have very long filaments
to place the anthers out where they can pick up the pollen
Figure 12.24 Acer saccharinum female and male flowers grains from the wind. In both flowers, petals and nectar
disks are absent (Figures 12.23 and 12.24).

228 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruit a schizocarp that breaks into samaras (thus


SAMAROID SCHIZOCARP fruits)

Acer spp. (Maples)

As with the Apiaceae family, Aceraceae produces


SCHIZOCARPS. However, these schizocarps are special in
that their MERICARPS are SAMARAS, or winged achenes.
Thus, the proper name for the fruits are SAMAROID
SCHIZOCARPS.
Figure 12.25 Acer platanoides SAMAROID SCHIZOCARPS.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Deciduous trees
▪▪ Opposite leaf arrangement and palmately lobed leaf
margin

Acer spp. (Maples)

Learn the vegetative characteristics of these plants by


examining the fresh material and herbarium specimens
provided (Figure 12.26). Then sketch out a typical
Aceraceae species leaf in Box 5, making sure to label its
parts.

Figure 12.26 Acer rubrum leaves.

Box 5

Lab 12 - Rosaceae, Aceraceae and Apiaceae 229


APIACEAE (UMBELLIFERAE)

carrot family ♂
♀ or ♀ Ca5 Co5 A5
Eudicots: Asterid Clade G②

INFLORESCENCE TYPE

▪▪ Simple or compound umbels

Angelica atropurpurea L. (Purplestem Angelica)


Daucus carota L. (Queen Anne’s Lace)

Examine the compound umbels provided (Figure 12.27).


Is an umbel inflorescence considered determinant or
indeterminant? Also, what part usually present in most
inflorescence types is missing in umbel inflorescences?

Figure 12.27 Umbel inflorescence of Daucus carota.

FLORAL CHARACTERISTICS

▪▪ Small flowers with a syncarpous gynoecium composed of


2 connate carpels and an inferior ovary
▪▪ Has a STYLOPODIUM (an enlargement at the style’s base

Anethum graveolens L. (Dill)

To observe the floral morphology typical of Apiaceae, you


Figure 12.28 Anethum graveolens umbel inflorescence. will examine and dissect a A. graveolens flower.

1. Observe a flower. What is its symmetry?

2. CALYX: Try to find the sepals. If they are not obvious,


examine under magnification the shoulder of the
ovary between and slightly outside of the petals. The
small bumps or protuberances that you see there are
actually the very reduced sepals. In some flowers the
sepals are completely absent and in others, such as
Coriandrum, the sepals are obvious.

Figure 12.29 Umbel inflorescence of Pastinaca sativa 3. COROLLA: The petals making up the corolla have an
flowers. Flowers possess a STYLOPODIUM. unusual shape, being pocketed or deeply contoured

230 Integrative Biology 335: Plant Systematics


and partially enclosing the anthers in these pockets.

4. ANDROECIUM: How many stamens do you see?


Because each of the five stamens in these flowers
matures at different times, not all stamens may be
seen. In younger flowers, the stamens may still be
enclosed by the petals. Also, note how the filament
attaches near the summit of the ovary but under the
swollen bases of the style (called STYLOPODIUM).

Figure 12.30 Osmorhiza longistylis umbel inflorescence


5. GYNOECIUM: How many carpels fused to create the
containing male and female flowers. syncarpous gynoecium of this flower and how can you
tell? Is the ovary superior or inferior and what is the
insertion type for the flower?

Osmorhiza longistylis (Torr.) DC. (Longstyle


Sweetroot)

In this species, there are separate staminate and


carpellate flowers (Figure 12.30). If both flower genders
are located on the same plant, what is the plant condition?

Note the inferior ovary of the carpellate flowers and the


prominant STYLOPODIUM at the top of the ovary (Figure
12.31).
Figure 12.31 Close-up of Osmorhiza longistylis male
and female flowers.

FRUIT TYPE

▪▪ Fruit is a SCHIZOCARP made up of 2 MERICARPS joined


together by a CARPOPHORE
▪▪ Mericarps contain oil tubes

Anethum graveolens L. (Dill)


Apium graveolens L. (Celery)
Cuminum cyminum L. (Cumin)
Foeniculum vulgare Mill. (Fennel)
Aethusa cynapium L. (Fool’s Parsley)
Osmorhiza claytonia (Michx.) C.B. Clark. (Sweetroot)

Observe the Apiaceae SCHIZOCARP fruits provided. In


Figure 12.32 Schizocarps of Zizia aurea, or Golden mature fruits, each carpel of the ovary separates from
Alexanders.
a slender axis called a CARPOPHORE (Figure 12.32).
When the fruit splits apart, each carpel section is called
a MERICARP. What other families have we studied this
semester produce SCHIZOCARP fruits?

Lab 12 - Rosaceae, Aceraceae and Apiaceae 231


HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Herbs that possess hollow stems, are aromatic and


sometimes are deadly poisonous
▪▪ Leaves are mostly pinnately compound with a sheathing
leaf base and display alternate arrangement

Coriandrum sativum L. (Coriander)

Try to pick out some of the vegetative characteristics of


this family, especially those listed above, on the coriander
plant on display. Also, if there are flowers on the plant,
check out their distinctive sepals.

Anethum graveolens L. (Dill)


Daucus carota subsp. sativus (Carrot)

Cut a cross-section of the fresh dill (Figure 12.33) and


carrot stems available. What do you notice about them?
Now sketch the cross section in Box 6.
Figure 12.33 Pinnately compound leaves of Anethum
graveolens, or dill.

Box 6

232 Integrative Biology 335: Plant Systematics


LABORATORY 13
ARACEAE, ORCHIDACEAE, IRIDACEAE, LILIACEAE,
ARECACEAE AND POACEAE

PHYLOGENETIC RELATIONSHIPS

In a cladogram from Zomlefer (1994) showing


the relationships of traditional “dicotyledons”
and monocotyledons, the traditional “dicots”
are paraphyletic, while the monocots are
monophyletic. Therefore, the traditional
dicot versus monocot split of angiosperms is
invalid. Synapomorphies for the monocots
include parallel-veined leaves, embryos
with one cotyledon, sieve cell plastids with
cuneate protein bodies and DNA sequences.
Stems with scattered vascular bundles and
adventitious roots also characterize the
monocots, but these characters also occur in
the “dicot” family Nymphaeaceae. There may
be exceptions (such as some monocots having
pinnate to palmate leaves with obviously
reticulate ventation patterns, like the Araceae),
but these are reversals of character states.
Three-merous flowers and the herbaceous
habit, characters often used to circumscribe the
monocots, also occur in Nymphaeaceae and
relatives and in some magnoliids. The monocot
clade plus the “dicot” families Aristolochiaceae,
Piperaceae and Nymphaeaceae are known as
the paleoherbs. In the Zomlefer cladogram, these families are closely allied with the monocots. In the Judd et al. text,
these families arise in the Magnoliid clade and more basally within the tree (e.g., in the “basal families” group). The
monocots are divided into 11 orders. In today’s Lab you will see a diversity of monocot families, but we will emphasize
only six: Araceae (Alismatales), Orchidaceae and Iridaceae (Asparagales), Liliaceae (Order Lilales, or the petaloid
monocots), Arecaceae (Arecales), and Poaceae (Poales). The families of the orders Arecales, Poales, Commelinales
and Zingiberales comprise the Commelinid clade of the Monocots.

Your textbook treats Liliaceae in the strict sense (recognizing only 22 genera), whereas in this class we take a broad
view of the family (and recognize Liliaceae sensu lato). Those species with inferior ovaries, a connate perianth with
adnate stamens, or a corona, or those with a “tree-like” or succulent habit are treated in at least nine other families
(see Table 19 in Zomlefer).

Poaceae is a large and complex family has been divided into 2-12 subfamilies and as many as 60 tribes. This lab is
meant to give you a broad overview of Poaceae morphology without going into depth on the subfamily characteristics.

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 233


ARACEAE
arum family ♂

♀ or ♀ Ca0 or 2-3 Co 0 or 2-3 A4-10 G 2 - 4
Monocot Clade

FLORAL CHARACTERISTICS

▪▪ Inflorescence is a SPADIX subtended by a SPATHE


▪▪ Flowers very reduced, often sunken into SPADIX

Zantedeschia spp. (Calla Lily)

Examine the SPADIX inflorescence, where many reduced


flowers are crowded onto a fleshy axis. Subtending the
SPADIX is a (often) colorful bract called a SPATHE. The tiny,
highly reduced flowers lack or have an inconspicuous
perianth, are often sunken into the SPADIX and can be
perfect or imperfect. When imperfect, female flowers are
generally located on the bottom of the SPADIX while male
flowers are located at the top.

Aglaconema modestum Schott (Chinese Evergreen)


Arisaema triphyllum (L.) Schott (Jack-in-the-Pulpit)

A. modestum contains both male and female flowers on its


spadix inflorescence (Figure 13.1). Thus, what is the plant
condition?
Figure 13.1 Close-up of an Aglaconema modestum
SPADIX and SPATHE.

In the Arisaema genus, the SPADIX has a large sterile


appendix and the flowers are confined to the lowermost
part (Figure 13.2). Further, male and female flowers are
found on separate spadix inflorescences. What is the plant
condition of A. triphyllum?

Figure 13.2 Arisaema triphyllum inflorescences.

234 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Produce multiple fruits of berries

Arisaema dracontium (L.) Schott (Green Dragon)


Arisaema triphyllum (L.) Schott (Jack-in-the-Pulpit)

Check out the herbarium specimens of Araceae. Each of


the flowers clustered on the SPADIX of these plants has
produced a berry fruit (Figure 13.3). What is the name
of a fruit formed from several separate flowers crowded
together on a compact inflorescence?

Figure 13.3 Arisaema triphyllum multiple fruit.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Perennial herbs or woody vines


▪▪ Basal leaves with sheathing bases
▪▪ Parallel, pinnate or palmate leaf venation
▪▪ Leaves often with RAPHIDES (needle-like crystals) of
calcium oxalate

Anthurium spp. (Flamingo Flower)

Check out the SPADIX inflorescence of Anthurium


Figure 13.4 An Anthurium SPADIX and SPATHE.
surrounded by its bright red SPATHE (Figure 13.4).
Like many other Araceae species, Anthurium has an
herbaceous habit. Additionally, its tissue contains bundles
of needle-like crystals (RAPHIDES) of calcium oxalate that
can cause painful injury to the mouth and throat. No matter
what your TA says, do not chew on these plants! What
type of leaf arrangement and venation does Anthurium
possess?

Amorphophallus titanum (Becc.) Becc. ex Arcang

Remember this plant from Lab 2: Vegetative Terminology?


The titan arum develops from a CORM and then spends
years in the vegetative part of its lifecycle (Figure 13.5).
What is a CORM?

After the vegetative stage is complete, the species will


bloom and emit a “rotting-fish-with-burnt-sugar” odor to
attract its natural pollinators, carrion beetles and flesh flies.
The titan arum is often said to have the “world’s largest
flower”. Based on what you now know about Araceae
species, is this a botanically accurate statement? Why or
why not?
Figure 13.5 An Amorphophallus titanum inflorescence.

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 235


ORCHIDACEAE
orchid family ♂ Ca3 CoZ 3 A1 or (2) (3)

G③
Monocot Clade

FLORAL CHARACTERISTICS

▪▪ Zygomorphic, resupinate flowers


▪▪ Stigma, style and androecium adnate forming a COLUMN

Dendrobium spp. (Dendrobium)


Phalaenopsis spp. (Moth Orchids)

The Orchidaceae family is characterized as having


strikingly zygomorphic, 3-merous flowers with a highly-
specialized middle petal (called a LIP or LABELLUM, which
may be highly modified in shape and color). The stigma,
style and androecium are adnate, forming a COLUMN
(Figures 13.6 and 13.7). This COLUMN of Orchidaceae
is unique among monocots and is analogous to the
GYNOSTEGIUM found in Asclepiadaceae. One stamen
makes up the androecium, with its cap-like anther
separated from the receptive part of the stigma by the
ROSTELLUM (the central, sterile lobe of the stigma.) Pollen
is agglutinated into POLLINEA. Label these structures on
the illustration provided below. Then, take some time to
appreciate the complexity of orchid flowers especially
when you visit the Orchid room of the Plant Biology
Collections.
Figure 13.6 A Bletilla striata flower.

Figure 13.7 Close-up of a Bletilla striata column.

236 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruit is a loculicidal capsule

Miscellaneous Orchidaceae spp.

The seeds found in the loculicidal capsules of orchid


species are microscopic and often require a fungal
symbiont to germinate (Figure 13.8).

Figure 13.8 Close-up of Cypripedium parviflorum


seeds.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Possess a PSEUDOBULB and VELAMEN covering the


adventitious roots

Miscellaneous Orchidaceae spp.

In epiphytic species of Orchidaceae, which are largely


limited to the tropics, the leaf blade often arises from a
PSEUDOBULB, and the whitish, cord-like, adventitious
roots have a VELAMEN covering (Figure 13.9). What is the
purpose of the VELAMEN covering?

Figure 13.9 A Phalaenopsis orchid.

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 237


IRIDACEAE
iris family ♂ Ca3 Co3 A3

G③
Monocot Clade

FLORAL CHARACTERISTICS

▪▪ Petaloid sepals (thus, petals and sepals may both be


referred to as tepals)
▪▪ Style is petaloid

Iris spp. (Iris)

To learn the floral characteristics of Iridaceae, you will


examine and dissect an Iris flower.

1. Examine a flower (Figure 13.10). What is its floral


symmetry?
Figure 13.10 An Iris flower.

2. CALYX and COROLLA: Observe the 3-merous petaloid


perianth of this flower. The large TEPALS of the outer
perianth (called “falls”) are spreading and deflexed,
while those of the inner whorl (called “standards”) are
smaller and erect (Figure 13.11).

3. ANDROECIUM: How many stamens make up the


androecium? Do you detect any adnation or
connation?

4. GYNOECIUM: In Iris, the styles of the gynoecium are


flat, petaloid and colorful. Observe how the three style
Figure 13.11 Longitudinal section of an Iris cristata branches curve along the tepals, forming a protective
flower. covering over the stamens. A flap-like stigma is
situated on the underside of each style branch near
the top. What is the gynoecium and ovary type? In
Box 1 Box 1, sketch and label a cross section of the ovary.

238 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruit is a loculicidal capsule

Iris pseudacorus L. (Pale Yellow Iris)


Sisyrinchium albidum Raf. (White Blue-Eyed Grass)

Examine the loculicidal capsules on display (Figure 13.12).


Make sure that you can differentiate between a loculicidal
and a septicidal capsule! Remember that a loculicidal
capsule will split open between the septa into the locule
to release its seeds whereas a septicidal capsule will split
Figure 13.12 Close-up of an Iris foetidissima capsule. open along its septa.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Perennial herbs with bulbs, rhizomes or corms


▪▪ Leaves are 2-ranked (EQUITANT) and sheathing at base

Crocus spp. (Crocus)


Iris spp. (Iris)

Iridaceae is composed of many perennial species that


possess bulbs, rhizomes or corms (Figure 13.13).

Neomarica spp. (Walking Iris or Apostle Plant)

Look at the leaves of this plant (Figure 13.14).


Figure 13.13 A Crocus corm. Characteristic of Iridaceae, the leaves are described as
2-ranked or EQUITANT. What does this mean?

Figure 13.14 EQUITANT leaves of an Iris plant.

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 239


LILIACEAE ♂ Ca3 Co3 A6 G③

lily family
OR
♂ Ca3 Co3 A6

Monocot Clade
G③

FLORAL CHARACTERISTICS

▪▪ Petals and sepals look similar (TEPALS)


▪▪ 3-merous flowers
▪▪ Anthers VERSATILE

Lilium spp. (Lily)

To learn the floral characteristics of Liliaceae, you will


examine and dissect a Lilium flower (Figure 13.15).

1. Examine a flower. What is its floral symmetry?

Figure 13.15 Lilium 3-merous flower with versatile


anthers

2. CALYX and COROLLA: How many sepals and petals


are there? In this family the sepals and petals are
often hard to distinguish from one another. What is the
name assigned to them in this case?

3. ANDROECIUM: How many stamens make up the


androecium? Do you detect any adnation or
connation? In this family the anthers are VERSATILE.
What does this mean?

Figure 13.16 Close-up of a Lilium flower.


4. GYNOECIUM: The syncarpous gynoecium of this
flower is made up of how many connate carpels? Is
the ovary superior or inferior? Make a cross section
Box 2 through the center of the ovary, examine it with your
dissecting scope, and then sketch it in Box 2. What is
the insertion type and placentation type?

5. DRAWING PORTFOLIO: Draw a floral diagram of a


longitudinal section (L.S.) of Lilium spp. as well
as a cross section (X.S.) of its ovary, making sure
to include its floral formula. Follow the format as
instructed in Lab 3: Floral Terminology.
240 Integrative Biology 335: Plant Systematics
We will only focus on the floral morphology characteristic
of the majority of Liliaceae species, but there are some
A6 species that require a unique floral formula. As you
can see, these species have 3 sepals in the calyx, 3

♀ Ca3 Co3 G③ petals in the corolla and 6 epipetalous stamens in the
androecium. The perianth is connate with adnate stamens
(“epi-tepalous”). The flower’s syncarpous gynoecium is
Figure 13.17 Close-up of a Polygonatum flower. composed of three connate carpels with a superior ovary.

FRUIT TYPE

▪▪ Fruit is a capsule

Hemerocallis spp. (Day Lily)

One of the fruit types commonly produced by Liliaceae


species is a capsule (Figure 13.18). What type of
gynoecium do capsules always arise from? Specifically,
what capsule type is on display?

Figure 13.18 A Hemerocallis capsule.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Perennial herbs with bulbs, rhizomes or corms

Allium spp. (Onion)


Allium sativum L. (Garlic)
Lilium spp. (Lily)
Tulipa spp. (Tulip)

Like Iridaceae, many members of Liliaceae have


underground structures that enable the plants to be
Figure 13.19 Bulbs, corms and rhizomes from Liliaceae perennial (Figure 13.19). Onions, garlic, lilies and tulips all
species. produce bulbs. Corms and rhizomes are also found in the
Liliaceae. What characteristics can you use to differentiate
between these stem types?

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 241


ARECACEAE (PALMAE)
palm family ♂

♀ or ♀ Ca3 Co3 A6 G 3 or ③
Monocot Clade

FLORAL CHARACTERISTICS

▪▪ Small perfect or imperfect flowers


▪▪ Flowers usually 3-merous

Chamaedorea spp. (Palm)

The perfect or imperfect flowers characteristic of this family


are found in highly branched inflorescences subtended by
large, basal sheaths (bracts). When perfect, the flowers
themselves consist of 3 distinct sepals, 3 distinct petals, 6
stamens and an apocarpous or syncarpous gynoecium of
3 carpels. Fusion of the floral series varies considerably
among the genera in this family.

Cocos nucifera L. (Coconut)

Coconuts produce inflorescences of imperfect flowers


(Figure 13.20). Note that the staminate flowers are above
the few carpellate flowers. What is the plant condition
when both staminate and carpellate flowers are located on
Figure 13.20 Cocos nucifera inflorescences. the same plant?

Roystonea borinquena O.F. Cook (Puerto Rico royal


palm)

Observe the closeup photograph of Puerto Rico royal palm


flowers in bloom (Figure 13.21).

Figure 13.21 Roystonea borinquena flowers.

242 Integrative Biology 335: Plant Systematics


FRUIT TYPE

▪▪ Fruit is either a drupe or berry

Cocos nucifera L. (Coconut)

Most species within the Arecaceae produce drupe fruits


(or 1-seeded berries) exhibiting fleshy, fibrous or leathery
mesocarps (Figure 13.22). In the case of the coconut, the
husk (consisting of the mesocarp and exocarp), which
is usually removed before the rest of the fruit is sold in
markets, is very fibrous. What purpose do you think this
Figure 13.22 Cocos nucifera drupe. fibrous mesocarp serves?

The part of the coconut you buy in grocery stores is the


“pit” or endocarp. The edible portion of a coconut is the
seed within the endocarp, and it comprises both solid
“meat” and liquid “milk” endosperm.

Lodoicea maldivica (J.F. Gmel.) Pers.

Lodoicea maldivica, commonly called Coco-de-mer or


Double Coconut, is a dioecious species that produces
the world’s largest seed, weighing up to 38.7 pounds
(Figure 13.23). To produce its “double coconut” fruit, the
flowers mature over a six to seven year span. L. maldivica
is currently listed as endangered and is restricted to the
Seychelles island in the Indian Ocean. Neat isn’t it?
Figure 13.23 Lodoicea maldivica drupe.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Trees or shrubs with unbranched trunks but without


secondary growth
▪▪ Leaves pinnately or palmately divided or compound with
large basal sheaths

Areca catechu L. (Betal Palm)

The “woody” trunk of species in this family is composed


of many interconnected vascular bundles and fibers;
true wood, from a permanent vascular cambium, is not
produced. Also, the trunks are often covered with the
remains of overlapping leaf bases, which add structural
rigidity. The persistent petioles can sometimes act as
spines. Make sure to check out the additional Arecaceae
Figure 13.24 Unbranched trunk of Areca catechu.
family members found in the Conservatory.

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 243


POACEAE
grass family ♂

♀ or ♀ Ca2 Co 0 A3 G 2-3
Monocot Clade

FLORAL CHARACTERISTICS

▪▪ Inflorescence is a SPIKELET
▪▪ SPIKELET and FLORET terminology important

Avena spp. (Oats)


Triticum spp. (Wheat)

The SPIKELET inflorescences found in Poaceae are


subtended by 2 bracts (GLUMES) and arranged in various
secondary inflorescences (i.e. spikes, racemes or
panicles). The flowers themselves are reduced, subtended
by 2 bracts (PALEA and LEMMA), and are collectively called
the FLORET (Figure 13.25). The perianth is reduced to
scales (LODICULES), and there are usually 3 stamens in
the androecium. Two to three connate carpels (typically 3
carpels with 2 styles) make up the syncarpous gynoecium
in these flowers.
Figure 13.25 Spikelet of Avena fatua.

Be sure you understand the relationship among primary


Box 3
(first or lower) and secondary (second or upper) GLUMES,
PALEA, LEMMA, FLORET, flower, RACHILLA and SPIKELET.
Create your own drawing of a Poaceae spikelet in Box
3, making sure to label all the terms listed above. Use
the live material on display as well as the provided
photographs to help you.

244 Integrative Biology 335: Plant Systematics


Zea mays L. (Corn)

Perhaps the most familiar grass in Illinois today is Zea


mays, or corn. The flowers are imperfect and the plants
are monoecious. The staminate flowers are produced in
tassels at the top of the plant, while the female flowers
are born in “ears”. In light of this, what actually is the stuff
protruding out of an ear of corn that we commonly call
“corn silk”?

Figure 13.26 Tassels and ears of Zea mays.

FRUIT TYPE

▪▪ Fruit is a CARYOPSIS

Hordeum vulgare L. (Common Barley)


Secale cereale L. (Cereal Rye)
Triticum aestivum L. (Common Wheat)
Zea mays L. (Corn)

All members of the Poaceae family produce CARYOPSIS


fruits (Figure 13.27). This fruit type is basically an achene
where the pericarp is fused to the seed coat. How does
this differ from an achene?

Figure 13.27 CARYOPSIS fruits of rice, wheat and corn.

HABIT AND VEGETATIVE CHARACTERISTICS

▪▪ Annual or perennial herbs

Avena spp. (Oats)


Triticum spp. (Wheat)

What is the habit of Poaceae species? The plants in


Poaceae have rhizomes or stolons and possess TERETE
(round in cross section) stems (CULMS) with hollow
internodes. Observe the 2-ranked leaves. What type
of leaf attachment is being demonstrated? Notice that
a LIGULE is present at the junction of blade and sheath
(Figure 13.28).
Figure 13.28 LIGULE of a Poaceae species.

Lab 13 - Araceae, Orchidaceae, Iridaceae, Liliaceae, Arecaceae and Poaceae 245


246 Integrative Biology 335: Plant Systematics
Practice Lecture Exam 1

For each of the multiple choice questions below, circle the one correct response.

1. What is taxonomy?

a) At the same time, the most basic and the most derived or synthetic field of biology.
b) The science of classification, especially the classification of biological organisms.
c) The study and description of the variation of organisms, the investigation of the causes and
consequences of this variation, and the manipulation of the data to produce a system of
classification.
d) All of the above.

2. Which of the following groups of taxa is written in the correct DESCENDING order of the taxonomic
hierarchy?

a) Magnoliopsida, Magnoliophyta, Magnoliales, Magnoliaceae, Magnolia virginiana


b) Magnoliophyta, Liliopsida, Poaceae, Cyperaceae, Zea mays
c) Magnoliophyta, Magnoliopsida, Asteraceae, Asterales, Aster tenuifolius
d) Magnoliopsida, Malvales, Sterculiaceae, Theobroma, Theobroma cacao

3. What is wrong with this sentence, paraphrased from a popular nature magazine? “There is only one
specie of plant, of which I am aware, that can be used in the religious exercises of the Buibui tribe.”

a) The name of the plant family is not provided, and it should be.
b) The word “specie” must be underlined.
c) The word “specie” is written incorrectly. “Species” is both singular and plural.
d) There are actually two species of plants that are used by the Buibuies.

4. Approximately how many extant species of flowering plants are there?

a) 520,000
b) 280,000
c) 130,000
d) 36,000

5. You discover a new species of Theobroma (T. cacao is the source of cacao, or chocolate) and want
to honor one of your first plant taxonomy instructors, Danielle Ruffatto, by naming the species after
her. Unfortunately, while you were collecting these plants you were hit on the head with a liana and
had to be told that your surname is Smith. What is the best correct scientific name of your new
species, student Smith?

a) Theobroma smithii Ruffatto
b) Theobroma cacao (Ruffatto) Smith
c) Theobroma cacao ‘Ruffatto’
d) Theobroma ruffattiana Smith

Practice Lecture Exam 1 247


6. A herbaceous plant:

a) always completes its life cycle in one growing season.
b) may have underground perennial structures like corms or tubers.
c) develops persistent above ground woody tissue.
d) All of the above.

7. Fragaria virginiana is the wild strawberry of eastern North America. Fragaria chiloensis is the wild
strawberry of western North America and South America. The cultivated strawberry is a hybrid of
these two species. The scientific name of this hybrid may be written correctly in what way?

a) Fragaria chiloensis + F. virginiana
b) Fragaria virginiana cv. ‘Sweet Bite’
c) Fragaria virginiana subsp. chiloensis
d) Fragaria × ananassa

8. The three plant genomes differ dramatically in size and this severely affects their utility in molecular
systematic study. Which genome(s) is(are) the smallest?

a) Chloroplast
b) Mitochondrion
c) Nucleus
d) Both plant mitochondrial and nuclear genomes are equally small in size.

Use this floral formula to answer the next THREE


questions on this page. This formula is representative
of all flowers on a single herbaceous plant.

9. The insertion of floral parts in this flower is:



a) hypogynous
b) epigynous
c) perigynous
d) synoecious

10. What type of gynoecium is exhibited by this flower?

a) monocarpous
b) apocarpous
c) syncarpous
d) perigynous

11. What type of placentation would you expect NOT to occur in this flower?

a) parietal
b) free-central
c) marginal
d) axile

248 Integrative Biology 335: Plant Systematics


12. Using the data matrix below and the method of maximum parsimony, which tree is produced from the
analysis of these data? For all characters, consider state “1” as the derived condition.

Species A 000101010
Species B 010000000
Species C 000100011
Species D 000010010

a) tree “a”
b) tree “b”
c) tree “c”
d) tree “d”

13. Consider tree “b” above only. If species C, D and B have been traditionally classified in the family
Urticaceae and species A in the family Moraceae, the family Urticaceae is:

a) monophyletic
b) paraphyletic
c) polyphyletic
d) synapomorphic

14. Again, consider tree “b” above and the family designations of question 13. What changes to the
traditional classification would have to be made in order to produce a classification that is truly
phylogenetic?

a) Transfer species A into the Urticaceae.
b) Transfer species B into the Moraceae.
c) Transfer species C into a new, yet to be described family.
d) No change to the traditional classification system would be required.

15. What family am I? I have flowers with 2 cauducous sepals, crumpled petals in bud, many stamens
arranged in whorls, milky or colored latex, and a syncarpous gynoecium.

a) Magnoliaceae
b) Ranunculaceae
c) Papaveraceae
d) Moraceae

16. What is unusual about the androecium of Hamamelis (Hamamelidaceae)?



a) The pollen is ejected forcibly through a ballistic propulsion mechanism.
b) The anthers are colorful in autumn.
c) The stamens are pendulous and serve to attract pollinators.
d) Staminodes are present and the anthers open by flaps.

Practice Lecture Exam 1 249


17. What family is illustrated below?

a) Magnoliaceae
b) Ranunculaceae
c) Papaveraceae
d) Hamamelidaceae

18. What family is illustrated below?

a) Hamamelidaceae
b) Magnoliaceae
c) Fagaceae
d) Betulaceae

250 Integrative Biology 335: Plant Systematics


19. What is a follicle?

a) Any dry, dehiscent fruit derived from a syncarpous gynoecium.


b) A dry fruit derived from a single carpel that opens along a single suture.
c) A small anatomical cavity or depression in which the ovule lies.
d) The product of the fusion of the egg and sperm nuclei.

20. Major structural rearrangements of which genome are known to be rare enough in evolution that
they can be used to demarcate major groups of plants and infer evolutionary relationships? Such
rearrangements include inversions and the gains and losses of genes or their introns.

a) Chloroplast
b) Mitochondrion
c) Nucleus
d) Both chloroplast and mitochondrial genomes are equally rarely rearranged.

21. What molecular technique involves the automatic replication of DNA using an enzyme and repeated
cooling and heating so that millions of copies of DNA are produced from one copy of DNA in a very
short period of time?

a) polymerase chain reaction (PCR)
b) restriction site mapping
c) DNA polymerase
d) amino acid translation

22. Which of the following shows an incorrect pairings between a character and its character states?

a) life span: annual, biennial, perennial
b) habit: herb, shrub, tree
c) root type: rhizome, stolon, corm
d) condition: synoecious, monoecious, dioecious

23. What is the best definition of a raceme?



a) A determinate compound (multi-branched) inflorescence
b) An indeterminate inflorescence in which all flowers have pedicels of equal length that arise from a
single region at the apex of the inflorescence axis
c) A compact inflorescence composed of a very short axis and usually sessile flowers
d) A simple, indeterminate inflorescence with an elongate single axis bearing pedicellate flowers

24. Using a strict botanical definition, what are the following: green beans, tomatoes, eggplants, corn,
and cucumbers?

a) vegetables
b) fruits
c) accessory fruits
d) seeds

Practice Lecture Exam 1 251


25. What type of fruit could possibly develop from a flower having a syncarpous gynoecium?

a) follicle
b) aggregate
c) capsule
d) all of the above

26. Compare and contrast the families Fagaceae and Betulaceae. For full points, you need to indicate
two shared characters and two characters where they differ. Use only those diagnostic floral/fruit
and inflorescence characters emphasized in lecture. [4 points]

2 shared chars.: _____________________________ and ______________________________

Fagaceae unique char.: __________________ Betulaceae unique char.: _________________

27. In general terms, outline the fig wasp–fig life cycle. [4 points]

28. What is a carpel? For full points, you must provide two different components to your answer. [2
points]

29. What is the difference between an aggregate fruit and a multiple fruit? For full points, you must
explain the difference clearly and provide one example of each. [4 points]

252 Integrative Biology 335: Plant Systematics


30. Use simple and neat drawings to illustrate each of the following terms. To obtain the point for each
drawing, you must show all necessary critical features and label as necessary to avoid any ambiguity.
[6 points; no partial credit (any error will receive a score of 0)]

A palmately compound leaf A lanceolate leaf with serrate margin

A X.S. of a syncarpous gynoecium Any leaf with a cordate base, an


with axile placentation acuminate apex, & palmate venation

A L.S. of any perfect, incomplete


flower A drupe X.S.

Practice Lecture Exam 1 253


31. The generic name for redbuds in Cercis. Linnaeus gave the specific epithet of canadensis to the
native redbud that occurs in Illinois. Most redbuds have dark pink/mauve flowers. In Missouri, a
few trees occur in the wild that have white flowers. Rehder described these white-flowered plants
as a botanical form, using the epithet of alba. What is the complete scientific name of the white-
flowered redbud? [2 points]

32. Draw a simple cladogram showing the relationships among the four major angiosperm groups:
eudicots, monocots, magnoliids, and the ANITA grade. [3 points]

BONUS: Why is systematics important in biological investigations and to society? [3 points]

TOTAL SCORE (out of 50): _______________ (maximum score with BONUS is 53 points)

254 Integrative Biology 335: Plant Systematics


PRACTICE LECTURE EXAM 2

Use this floral formula to answer the first four questions on


this page. This formula is representative of all flowers on a
single herbaceous plant.

1. The insertion of floral parts in this flower is:



a) hypogynous
b) epigynous
c) perigynous
d) synoecious

2. What family is best characterized by this floral formula?

a) Brassicaceae
b) Oleaceae
c) Ericaceae
d) Onagraceae

3. Which of the following statements is CORRECT given the floral formula above?

a) The stamens are epipetalous.


b) The flower is imperfect.
c) The flower is wind pollinated.
d) The flower is actinomorphic.

4. What type of fruit(s) could possibly develop from this flower?

a) follicle
b) capsule or berry
c) aggregate
d) silique or silicle

5. While dissecting a flower, you observe that it has a syncarpous gynoecium with parietal placentation,
2 locules, 1 style, 1 stigma, tetradynamous stamens, and 4 sepals and 4 petals arranged in a cross-
like formation. How many carpels have likely fused to produce the gynoecium of this flower?

a) one
b) two
c) four
d) The data are insufficient to determine this.

Practice Lecture Exam 2 255


6. Sporophytic self-incompatibility:

a) is controlled by multiple alleles of a single gene


b) ensures that a plant cannot produce a zygote with its own pollen
c) is determined by the interaction between the pollen exine and stigma tissues
d) all of the above

7. In the Indian balsam (Impatiens sp.), the nectar is held deep in the flower so that the bee has to enter
the flower to get it. When all the pollen is removed, the stamens fall off to expose the sticky, receptive
stigma. What type of breeding system is exhibited by this species?

a) protogyny
b) protandry
c) polygamodioecy
d) heterostyly

8. A legume fruit is derived from what type of gynoecium?

a) monadelphous
b) apocarpous
c) syncarpous
d) monocarpous

9. Fill in the blanks. In a species exhibiting a heterostylous breeding system, pollen from a ________
flower must reach the stigma of a ________ flower in order for fertilization to be successful.

a) pin, thrum
b) pin, pin
c) staminate, carpellate
d) thrum, thrum

10. Members of Fabaceae subfamilies Faboideae and Caesalpinioideae are alike in that they share:

a) diadelphous stamens
b) zygomorphic corollas
c) a keel formed from the fusion of two petals
d) all of the above

11. Plants have evolved a variety of adaptations that can either increase, reduce or maintain the amount
of genetic heterozygosity in a population. Which of the following kinds of breeding systems actually
increases the fitness or evolutionary potential of a species?

a) self-pollination
b) apomixis or agamospermy
c) gametophytic self-incompatibility
d) cleistogamy

256 Integrative Biology 335: Plant Systematics


12. Which of the following androecium modifications is an INCORRECT match?

a) monadelphous stamens Malvaceae


b) tetradynamous stamens Brassicaceae
c) diadelphous stamens Fabaceae subfamily Caesalpinioideae
d) stamens arranged in catkins Salicaceae

13. As seen in the video “Branching Out,” the acacias (Fabaceae subfamily Mimosoideae) of Africa have
relied on elephants to:

a) trample and bury their seeds during the long, hot, dry seasons so that they don’t dry out before
they are ready to germinate
b) disinfect the grub-infested fruits by ingesting and defecating the seeds
c) facilitate pollination of its flowers by knocking against or rubbing the trees
d) disperse its fruits due to their claw-like appendages which attach harmlessly to the thick elephant
skin

14. The tropical durian fruit, with its foul-smelling rind that wafts over great distances and its sweet
caramel custard-like pulp (... yeah, really it’s disgusting!), is favored by what type(s) of animal?

a) rhinos
b) orangutans and tigers
c) cassowaries
d) gnat flies

15. What family am I? In temperate regions, I am a perennial herb. My flowers are zygomorphic with
prominent nectar guides. One of my anterior petals has a large spur, and two of my lowermost
anthers bear gland-like appendages (nectaries) which extend into this spur. When an insect (usually
a bee) probes for nectar, it gets showered by pollen.

a) Brassicaceae
b) Violaceae
c) Ericaceae
d) Asclepiadaceae

16. What is a diaspore?

a) Any unit of dispersal, no matter what it is morphologically


b) The smallest of all spores, produced by fungi and dispersed most widely
c) The breaking up and scattering of plant vegetative tissue into new areas
d) Spores produced by a puffball and knocked from it like puffs of smoke

Practice Lecture Exam 2 257


17. What family am I? I have yellow, sympetalous, imperfect flowers, my anthers are often connate, and
my ovary is inferior. I am a herbaceous vine with tendrils. My fruit has a hard, leathery rind, its inside
is fleshy with numerous seeds, are there are no septa. And, I am absolutely terrible baked into a pie.

a) Brassicaceae
b) Malvaceae
c) Oleaceae
d) Cucurbitaceae

18. Hamamelis (witch hazel), dwarf mistletoes, Himalayan balsam (Impatiens sp.) and squirting cucumber
(Echbalium) share what kind of seed dispersal mechanism?

a) shaker
b) hydroscopic
c) adhesion by viscin threads
d) ballistic

Use this phylogeny of four families


with mapped character states (A-G) to
answer the following three questions.

19. The character state at position E is most likely what?

a) The presence of betalain pigments


b) The loss of the intron in chloroplast gene rpl2
c) The presence of a gynophore
d) All of the above

20. Which character states are mapped at the INCORRECT position?



Position Character State(s)
a) A Monadelphous stamens, palmately lobed and veined leaves
b) B Two persistent sepals, pyxis fruit
c) D Succulent leaves or succulent stems
d) G Apetalous flowers, ochrea, sepals often in two whorls of three

21. At what position on the tree would you expect to find the following synapomorphies: opposite, simple
leaves; swollen nodes; distinct clawed petals; capsular fruit with apical teeth; cymose inflorescence?

a) A
b) C
c) E
d) G

258 Integrative Biology 335: Plant Systematics


22. Which of the following statements is TRUE?

a) Champaign-Urbana receives, on average, less rain per year than London, England
b) About ten percent (by area) of the original prairie remains in Illinois today
c) Illinois is part of the “mixed-grass” prairie region
d) The word prairie comes from the French for “a meadow grazed by cattle”

23. Prairies formed in Illinois when its climate became substantially warmer and drier (and within the
relatively short time of 500 to 800 years, most of the forests in Illinois died out). This significant
warming – called the hypsithermal interval – occurred how many years ago?

a) 100,000
b) 12,000
c) 8,300
d) 1,000

24. Because North American prairie ecosystems are recently developed, there are actually very few plant
species that are unique (endemic) to them. Where, then, did the species we find on Illinois prairies
come from?

a) They migrated from cooler regions in the north


b) They migrated from regions in the southeast
c) They migrated from the eastern deciduous forests
d) All of the above

25. What family is illustrated below?

a) Euphorbiaceae
b) Caryophyllaceae
c) Asclepiadaceae
d) Brassicaceae

Practice Lecture Exam 2 259


26. Plants of the Cactaceae, Euphorbiaceae and Asclepiadaceae families, with their succulent stems
bearing spines, may look superficially similar when growing in arid regions. Provide two diagnostic
floral or vegetative features for each family that could be used to differentiate them from the other two
families. Each feature you choose should be unique to that family. [6 points]

Cactaceae: 1) ______________________________

2) ______________________________

Euphorbiaceae: 1) ______________________________

2) ______________________________

Asclepiadaceae: 1) ______________________________

2) ______________________________

27. Congratulations! You have just been appointed assistant curator of the Alfred Rehder Garden of
beans and peas at the Chicago Botanic Garden. Unfortunately, to create this garden, they had
to destroy a small plot of tomatoes. Your first job is to construct a proper dichotomous key for the
identification of the three subfamilies of Fabaceae: Mimosoideae, Caesalpinioideae, and Faboideae.
Your first couplet, however, should distinguish between tomatoes (Solanaceae) and all legumes
(Fabaceae). [4 points]

28. Select the most appropriate mode of pollination for each of the floral features described. Fill in the
blank with the letter of the best match. There is only one correct answer per question, and each type
of pollinator is only to be used once. (4 points)

a) hummingbirds
b) bats
c) beetles
d) bees
e) carrion beetles and flies
f) bush mice and other small mammals
g) butterflies
h) hawkmoths

_____ Purple or brown-colored actinomorphic flowers sometimes smelling strongly of rotten meat, and
no nectar or other award offered (examples, Rafflesia, titan arum, skunk cabbage, pawpaw)

260 Integrative Biology 335: Plant Systematics


_____ Large, sturdy, bowl-shaped flowers opening at night, white or cream in color, strong odor smelling
musty or like fermenting yeast, abundant nectar and pollen, and no nectar guides (examples,
saguaro cactus, durian tree)

_____ Sturdy flowers accessible near the ground, usually brown in color or sometimes white or creamy,
opening at night with a yeasty odor, and with abundant nectar and/or pollen (example, Protea)

_____ Actinomorphic, stiff, wide tubular, and easily accessible flowers with hanging stamens opening
during the day, Ferrari-red in color in the New World, no obvious odor, no nectar guides, but
loaded with abundant nectar (examples, Fuchsia, Aquilegia and Lychnis)

_____ Actinomorphic flowers open during the day with narrow tubular corollas, erect anthers, and a wide
landing platform, yellow, blue, or pink in color, often with nectar guides and abundant nectar
hidden deep in spurs (examples, soapwort, Phlox, many Asteraceae)

_____ Bright yellow, blue or white zygomorphic flowers opening during the day with sweet odor,
abundant nectar, nectar guides, and sometimes reflecting UV light in a “bulls-eye” pattern
(examples, many Fabaceae, Lamiaceae, and Scrophulariaceae, Salix)

_____ Actinomorphic white to sometimes pale green flowers with narrow tubular corollas, opening at
night or dusk, with a heavy sweet odor at night, abundant hidden nectar, and no nectar guides
(examples, Angraecum orchid, honey suckle)

_____ Large, bowl-shaped, unspecialized actinomorphic flowers with numerous floral parts and ovules
protected on an elongate receptacle or in a hypanthium, dull, creamy-white or green in color,
strong fruity or aminoid odor, no nectar guides, and abundant pollen and sometimes nectar
(examples, strawberry, Magnolia, water lily)

29. Draw the following. There is no need to label your drawings. [2 points]

A L.S. through a flower from any A single male flower of Euphorbia


typical member of the asterid clade (Euphorbiaceae)

30. For each of the following figures from your text, answer the following in the spaces to the right of
each figure: (a) Family or subfamily? (b) List two diagnostic floral/fruiting features for that taxon
that can be seen in the illustration. (c) Indicate the major clade to which the taxon belongs (i.e.,
asterid, Caryophyllales, rosid). [3 points for each figure; 9 points total]

Practice Lecture Exam 2 261



a) ________________________

b) ________________________

c) ________________________


a) ________________________

b) ________________________

c) ________________________


a) ________________________

b) ________________________

c) ________________________

262 Integrative Biology 335: Plant Systematics


BONUS (5 points)

List four reasons why seed/fruit dispersal is important to a plant species. [4 points]

1)

2)

3)

4)

Name one plant whose seed/fruit is dispersed by water. [1 point]

Total Score: __________________ (maximum score, with bonus, is 55 out of 50)

Practice Lecture Exam 2 263


264 Integrative Biology 335: Plant Systematics
THE STORY OF BLACKEYED SUE

I rose early, at four o’clock, the morning glory still iris away. I was worried. Anemone of mine, Johnny
Jump Up, was looking for me, and I’d heard he was carrying a pistil, a 357 magnolia. I ironed a
periwinkled blouse, got dressed, and took a sprig of a dusty Miller’s beer. Johnny Jump Up. He was
one of several rhizomes who’d gone to seed in Forsythia, Montana. He was convicted of graft in 1984,
arrested again in ‘85 for digging in coreopsis. Then he drifted on the wind up to my neighborhood, the
corner of Hollyhock and Vine. He was a petal pusher in a phloxhous nearby.

I knew he was trouble when he rode-a-dendron to my house and said, “Hey, little Black-eyed Susan,
wanna come over to my place and take a look at my vetches?” I didn’t want to tell him that in all the
cosmos, there was no one for me but Sweet William, so I said no, I was taking care of a pet dogwood
that’d had a litter of poppies, which was weird ‘cause she’d just been spaded. But Johnny had no sense
of humus. He stamped his foot with impatiens. “You’ll rue the day you turned me down,” he snapped.
Then he spit a wad of salvia into the petunia on my portulaca and stalked away. “Forget me not, Sue,
‘cause I’ll be zinnia.”

Ever since then, he’d cultivated a relationship with Lily of the Valley, a self-sowing biennial. One day, I
aster what she seed in him. “Mum’s the word on this,” she said. “He’s got a trillium dollars in the bank.”

“A trillium?” I snorted. “He’s lime to you. Besides, what about love?”

“Alyssum,” she said. “You bleeding hearts are all alike. Kid, you can go for a guy who’ll azalea with
affection. Orchid, you can be like me and try to marigold. Now begonia.”

Now I was in my kitchen mullein over these past events. But it was thyme to quit dilly-dahliaing. The
calendula read August 3rd, and Johnny had sworn to propagate vengeance before the snowdrop.

I hopped into my autolobelia and drove over to Daisy’s for help. Daisy was a pretty little transplant from
Florida, who’d wilted in the humidity there, but was rooted in the well-drained soil of Bloom County. She
mostly took care of her babies breath, but lately she’d branched out and was columbining work with home
life. “We’re all sick today, I think it’s gaillardia,” she said. “Even the cat’s got harebells. If we could take a
knapweed be OK.” Her face was a blight yellow. She’d be no help.

I beetled feet over to Sweet William’s. “Will, am I gladiolus to see you!”

“And Blackeyed Sue. I been prayingmantis see you. Let’s lilac in the snow on the mountain before it all
melts down the geranium. Let’s ride a sage to Tansynia. It’s only a chamomile away.”

“Don’t be fritillary, honeysuckle,” I said, clinging to him. “Look. Here comes the clematis of the story.”

Uh oh. Johnny had hired Pete Moss, a bearded iris-man to do me in. He was wearing a blue nectar and
larkspurs. He had larva men with him. The pests. They began to charge. In all the confuscia, I said to
Will, “Stem still and give me some ground cover.” I ran down the primrose path in my ladyslippers right
towards Pete.

The Story of Blackeyed Sue 265


“Don’t gimme any flax, bud, or I’ll slug ya,” I said.

“You’ll look dandelion in the alley.” “Don’t gimme any flax bud,” Pete quoted me verbena. It nettled me. I
clovered him with a 2 x 4.

“Sound the timpansy,” we sang. “We won!”

“Curses,” moaned Pete, “foliaged again.”

I noticed Johnny Jump Up planted on the border. “I’ve sunk pretty loam, Sue, but now I’m turning over a
new leaf” he said.

“Bouquet,” I said. And he did. And Will and I lived pearly everlasting.

266 Integrative Biology 335: Plant Systematics

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