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Marcescence in oaks
(David Remmen)
Subclass Rosidae
Monocots
Basal Caryophyllids Rosids
Eudicots
Eudicots
Asterids
Subclass Rosidae
Diverse group: > 15 orders, >100 families, >50,000 species
Major Rosidae families in Montana
Rosids I
Rosaceae (roses)
Salicaceae (willows)
Violaceae (violets)
Fabaceae (peas)
Betulaceae (birches)
Rosids II
Brassicaceae (mustards)
“Aceraceae” (maples)
Myrtales
Onagraceae (evening primroses)
Saxifragales
Saxifragaceae (saxifrages)
Grossulariaceae (gooseberries)
Other Rosids I
Other Rosids II
Rosaceae - Rose family
Distribution: world-wide; primarily northern
hemisphere => 100 genera, 3000 species

In Montana: 8th largest; 31 genera, 85 species
Economic plants/products: roses, apples, pears,
cherries, plums/prunes, almonds, peaches, apricots,
strawberries, raspberries, blackberries, many
ornamental plants
Rosaceae - Rose family
Life history/habit
herbs, shrubs or trees (many woody)
Leaves
alternate,
simple to compound
usually dentate (w/ glandular teeth)
often w/ paired stipules at leaf-base


Rosaceae - Rose family
Inflorescences
various, usually indeterminate (racemes etc.)
Flowers
radially symmetric
mostly bisexual (perfect)
with hypanthium (floral cup)
Rosa
Prunus

Rosaceae - Rose family
Sepals
5
distinct above
Petals
fused floral cup
5
(hypanthium)
Stamens
10 - 
Gynoecium
extremely variable!
Fruit types
follicle, achene
drupe, pome, or hip
aggregate of achenes or drupes
Rosa
Prunus
Prunus
Prunus
Rosaceae - Rose family
Amelanchier
Rosa
Potentilla
Rosaceae - Rose family
Variation in gynoecium and fruits
Pyrus (apple/pear)
Rubus, Fragaria, Potentilla
(raspberry and strawberry)
Rosa (rose)
Prunus (cherry)

Rosaceae - Rose family
Spiraea-type gynoecium
superior ovary, 2-5 unfused carpels => follicles
Rosaceae - Rose family
Spiraea-type gynoecium -- Physocarpus (Ninebark)
Rosa - Roses
Rosa-type gynoecium
half-inferior ovary
swelling of hypanthium => fleshy hip
many unfused carpels => achenes
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Rosa - Roses
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Rosa - Roses
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Rosa - Roses
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Rose “thorns” are technically prickles = extension of shoot epidermis
Rosaceae - Rose family
Pyrus-type gynoecium
inferior ovary of 2-5 fused carpels
swelling of hypanthium => fleshy pome
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Amelanchier - Saskatoon, Serviceberry
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Sorbus - Mountain Ash
Crataegus - Hawthorn
Rosaceae - Rose family
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Prunus-type gynoecium
superior ovary of 1 carpel
swelling of ovary => fleshy drupe w/ hard seed-covering
Prunus - Cherry
P. virginiana (Choke cherry)

P. pennsylvanica (Pin Cherry) and others
Rosaceae - Rose family
Fragaria-type gynoecium
superior ovaries, many unfused carpels
swelling of receptacle => accessory fruit w/ achenes
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Rubus - Raspberries, Blackberries
aggregate fruit of
many small drupes
Potentilla - Cinquefoils
Potentilla recta - Sulfur cinquefoil

Pentaphylloides - Shrubby Cinquefoil
Geum triflorum - Prairie Smoke
Saxifragaceae - Saxifrage family
Distribution
north temperate and boreal zones, diverse habitats
In Montana
14 genera
~50 species
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Saxifragaceae - Saxifrage family
Growth habit
primarily herbs, sometimes succulent
Leaves
alternate; often basal
simple, often palmately lobed

Saxifragaceae - Saxifrage family
Inflorescences
racemes, panicles, corymbs
Flowers
bisexual
radially symmetric
with hypanthium

Saxifragaceae - Saxifrage family
Sepals
5
Petals
5
Stamens
3-10
hypanthium

Saxifragaceae - Saxifrage family
Gynoecium
2 fused carpels => 2 styles
ovary position variable (inferior or superior)

Saxifragaceae - Saxifrage family
Fruit type
capsule

Saxifragaceae - Saxifrage family
Heuchera
Alumroot
Saxifragaceae - Saxifrage family
Parnassia
Grass-of Parnassus
Saxifragaceae - Saxifrage family
Saxifraga
S. odontoloma
wet areas/woodlands
S. rhomboidea
open grasslands
S. bronchialis
dry, rocky areas
Saxifragaceae - Saxifrage family
Boykinia
Mitella
Lithophragma - Woodland star
Habit
perennial herbs
Leaves
basal rosette
palmately divided leaves
Inflorescence
short raceme

Lithophragma - Woodland star
Sepals
5, fused
Petals
5, each divided in 3
Stamens
10
Gynoecium
2 fused carpels = 1 ovary
partially inferior

Lithophragma - Woodland star
Lithophragma parviflorum
Coevolved w/ Greya moth
moth is primary pollinator
but also lays eggs on seeds
costs/benefits of interaction vary
across environments
- where Greya are only pollinator
=> mutualism (+/+)
- where other pollinators common
=> parasitism/defense

Viola - Violets
Type genus of the family Violaceae
Habit
perennial herbs
often with basal rosette
Leaves
often cordate (heart-shaped)
alternate or basal
w/ fringed stipules

Grossulariaceae - Gooseberry family
Only 1 genus worldwide (Ribes)
~12 species of Ribes here
Habit
shrubs
prickly (gooseberries)
or not (currants)
Ribes - Gooseberries, Currants
Leaves
palmately lobed
Flowers
similar to Saxifragaceae
ovary ± inferior
corolla sometimes fused

Ribes - Gooseberries, Currants

Ribes - Gooseberries, Currants
Fruit type = berry

Viola - Violets
Flowers
generally solitary from base or in small clusters
bilaterally symmetric
bisexual

Viola - Violets
Sepals
5, distinct
Petals
5, distinct
1 spurred
Stamens
10

Viola - Violets
Gynoecium
3 fused carpels
1 locule (no walls)
superior
Fruit type
capsule

Viola - Violets
Dispersal
Seeds with
arils = elaiosomes
Dispersed by ants =
myrmecochory

Viola - Violets

Infloresescence types
Indeterminate
Determinate
1
3
3
2
umbel
corymb
2
3
3
cyme
1
2
raceme
x
x
x
x
6
panicle (branched)
spike (no
pedicels)
3
x
5
4
helicoid cyme
Parts of an inflorescence
terminal bud/youngest flower
rachis
pedicel (flower stalk)
older flower becoming fruit
peduncle (inflorescence stalk)
Two broad categories of inflorescences
Indeterminate
flowering begins at base
Two broad categories of inflorescences
Indeterminate
flowering begins at base
Two broad categories of inflorescences
Indeterminate
flowering begins at base
Two broad categories of inflorescences
Indeterminate
flowering begins at base
Two broad categories of inflorescences
Indeterminate
flowering begins at base
Determinate
apical bud flowers first
Two broad categories of inflorescences
Indeterminate
flowering begins at base
Determinate
apical bud flowers first
Indeterminate
raceme
simple (unbranched)
stalked flowers
Epilobium
Indeterminate
corymb
umbel
raceme
panicle (branched)
spike (no pedicels)
Determinate
cyme =
branched
stalked flowers
bracts at branch points
3
1
3
2
3
2
3
Determinate inflorescences
helicoid cyme
- lateral buds on 1 side
- curls into spiral
- common in Boraginaceae
1
2
x
x
x
x
6
x
5
Amsinckia (Boraginaceae)
3
4