Poales II: wind pollinated families

Poaceae - the grasses

showy flowers, insect or bird pollinated +/- reduced flowers, insect or wind pollinated

the second large independent shift to reduced flowers and spikelets for wind pollination 4th largest family - 620 genera, 10,000 species most important family (ethnobotanically)

reduced flowers, wind pollinated

Poales II: wind pollinated families

Evolutionary trends: nectar to pollen gathering to wind pollination reduced flowers - loss of perianth unisexuality sometimes bracts become important flowers to florets in spikelets showy flowers, insect or bird pollinated +/- reduced flowers, insect or wind pollinated reduced flowers, wind pollinated

Poaceae - grasses
Poaceae related to more typical, although reduced, flowered graminoid monocots with 6 tepals Southern Hemisphere!

Poaceae - grasses
Poaceae related to more typical, although reduced, flowered graminoid monocots with 6 tepals

Early floral development in Streptochaeta, Elegia, and Joinvillea.

Streptochaeta Poaceae

Restionaceae

Joinvilleaceae

Whipple C J et al. PNAS 2007;104:1081-1086

Poaceae - grasses

Poaceae - grasses
2/4 2 / 12 2 / 12 genera / spp
BEP

has whole genome doubling been a key innovation for grass diversification? 10,000 vs. 30 spp.

658 / 10,000

PACCAD

Whole genome duplications in Angiosperms (Soltis et al. 2009, Amer J Bot)

Likely occurrence of whole genome duplication in grasses

Poaceae - grasses
node

Poaceae - grasses
Adaptive features intercalary meristem grazing & fire response silica in stems C4 photosynthesis

Vegetative features jointed, hollow, circular stems (culms) leaves 2-ranked or spiralled blade, sheath, and ligule intercalary meristem above nodes Nebraska grassland 25 mya

blade ligule sheath

Poaceae - grasses
Adaptive features first diversified ca. 70 mya in late Cretaceous tropical forest understories major radiations during formation of grasslands in mid Tertiary is this set of features the key innovation?

Poaceae - grasses
Defining feature of grasses are the spikelet and its florets

Strmberg C A E PNAS 2005;102:11980-11984

Poaceae - grasses
The main unit of the inflorescence is the spikelet which is composed of 2 glumes (spikelet bracts) and 1 or more florets

Poaceae - grasses
Each floret is surrounded by two floret bracts - the outer lemma and the inner palea (usually not seen until anthesis - when florets open)

Brome

Oats

Brome

Oats

spikelet glumes florets Dactylis glomerata Orchard grass

lemma palea Dactylis glomerata Orchard grass

Poaceae - grasses
This spikelet with two glumes has two florets each with two floret bracts - the outer lemma and the inner palea

Poaceae - grasses
Although considerable variation occurs in florets (among species or within a spikelet), most of our species have the following floret structure:

glumes lemma palea

Perianth represented by 2 lodicules

What is function of lodicules?

Poaceae - grasses
anthesis expose anthers & styles fungal endophytes preventing spores entering fruit?

Poaceae - grasses
fungal endophytes (ascomycetes) produce physiologically active alkaloids anti-herbivory defense against grazing mammals (defensive mutualism)?

What is function of lodicules?

Nebraska grassland 25 mya

Poaceae - grasses
panic grass from Yellowstone Natl Park geothermal tolerant >50oC requires fungus endophyte fungus requires a virus to confer this thermotolerance effect

Poaceae - grasses

phylogenetic analysis of bamboos turned out to be phylogeny of endophytic fungi!

Lynn Clark Jonathan Wendel

Poaceae - grasses
Although considerable variation occurs in florets (among species or within a spikelet), most of our species have the following floret structure:

Poaceae - grasses
Although considerable variation occurs in florets (among species or within a spikelet), most of our species have the following floret structure:

Toby Kellogg Univ Missouri SL Perianth represented by 2 lodicules Stamens 3 Superior gynoecium of 2 fused carpels One ovuled fruits called a grain or caryopsis = seed fused to ovary wall Perianth represented by 2 lodicules Stamens 3 Superior gynoecium of 2 fused carpels One ovuled fruits called a grain or caryopsis = seed fused to ovary wall

What parts homologous to other flowers?

What parts homologous to other flowers?

Poaceae - grasses
lodicules = petals palea/lemma = sepals

Poaceae - grasses

Grass phylogenetics
What parts homologous to other flowers?

Basal grasses

Poaceae - grasses
South American origin!

Poaceae - grasses
Streptochaete

BEP

PACCAD

Subfamily Anomochlooideae
(no spikelets, lodicules)

Poaceae - grasses
Anomochloa marantoidea

Poaceae - grasses
Subfamily Pharoideae
(herbaceous bamboos)

4 stamens!

Pharus

Poaceae - grasses
Bambusa

Poaceae - grasses

Grass phylogenetics
Core grasses
Ochlandra

Subfamily Bambusoideae
(6 stamens, 3 lodicules, 3 stigmas)

Poaceae - grasses

Poaceae - grasses

Subfamily Ehrhartoideae
(stamens 6, but 2 styles) Oryza sativa - rice 2nd most important crop plant in the world

Subfamily Ehrhartoideae
(stamens 6, but 2 styles) Ziziana aquatica - wild rice Important native American food; unisexual spikelets

Poaceae - grasses
Tribe Poaeae

Poaceae - grasses
Dactylis glomerata - orchard grass

Tribe Poaeae

Poa annua - bluegrass

Subfamily Pooideae
(Spikelets with more than one grain forming floret; Lemma with 5 nerves)

Subfamily Pooideae
(Spikelets with more than one grain forming floret; Lemma with 5 nerves)

Poa pratensis - Kentucky bluegrass

Bromus inermis - bromegrass

Poaceae - grasses
Tribe Poaeae

Poaceae - grasses
Subfamily Pooideae Tribe Agrostideae
(awned lemma)

Colobanthus quitensis Deschampsia antarctica Only 2 angiosperms - Antarctic hairgrass and a Caryophyllaceae occur on north facing slopes with mosses and liverworts

Antarctic Peninsula - 600 mi or 1000 km from South America

Ammophila breviligulata - marram grass

Poaceae - grasses
Subfamily Pooideae Tribe Agrostideae
(awned lemma)

Poaceae - grasses
Subfamily Pooideae Tribe Agrostideae
(awned lemma) Phalaris arundinacea Reed canary grass

Calamagrostis canadensis bluejoint grass

Avena sativa - oats

Invasive species of wetlands

Poaceae - grasses
Subfamily Pooideae Tribe Hordeae/Triticeae
(symmetrical spikes)

Poaceae - grasses
Subfamily Pooideae Tribe Hordeae/Triticeae
(symmetrical spikes)

Elymus hystrix [Hystris patula] bottlebrush

Elymus canadensis Wild rye Triticum aestivinum wheat Hexaplod food crop

Poaceae - grasses
Subfamily Pooideae Tribe Hordeae/Triticeae
(symmetrical spikes)

Poaceae - grasses
Subfamily Arundinoideae
(large inflorescences)

Phragmites australis - common reed Agropyron repens quackgrass Circumboreal species; nonnative populations have become invasive and displaced native populations

Elytrigia wheatgrass

Poaceae - grasses
Subfamily Arundinoideae
(large inflorescences)

Poaceae - grasses
Subfamily Aristidoideae
(Awns of lemma divided into 3 parts)

Cortaderia - plume grasses from pampas

Aristida tuberculosa - 3-awned grass

Poaceae - grasses
Subfamily Chloridoideae
(Spikelets arranged often one-sided)

Poaceae - grasses
Subfamily Chloridoideae
(Spikelets arranged often one-sided)

Spartina pectinata Prairie cord grass Eragrostis cilianensis Stinkgrass

Bouteloua curtipendula Sideoats grass

Sporolobus heterolepis - Prairie dropseed

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Tribe Paniceae

Tribe Paniceae

Digitaria - crabgrass Panicum virgatum - switchgrass Panicum sp. - panic grass Setaria - foxtail

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Tribe Andropogoneae
(spikelets paired on linear inflorescence)

Tribe Andropogoneae
(spikelets paired on linear inflorescence)

Andropogon gerardii - big bluestem

Tripsacum - eastern gammagrass

Andropogon - bluestem

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Tribe Andropogoneae
(spikelets paired on linear inflorescence)

Tribe Andropogoneae
(spikelets paired on linear inflorescence)

Sorghastrum nutans - Indian grass Saccharum sugarcane Sorghum sorghum

Poaceae - grasses
Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Poaceae - grasses
The origin of maize from teosinte wild relatives in Mexico involved few genes Subfamily Panicoideae
(2 florets, bottom reduced, sterile)

Tribe Andropogoneae
(spikelets paired on linear inflorescence)

Tribe Andropogoneae
(spikelets paired on linear inflorescence)

Female spikelets solitary but condensed into cob or spike

Male spikelets paired in panicle or tassel

Zea mays - maize

Zea mays - maize

Tripsacum - teosinte

John Doebley

grasses you collected!

Picture key to grasses!

Setaria spp. foxtail Andropogon gerardii Andropogon virginicus big bluestem little bluestem

Setaria spp. foxtail Andropogon gerardii Andropogon virginicus big bluestem little bluestem

Grasses of Iowa www.eeob.iastate.edu/research/iowagrasses

Sorghastrum nutans Indian grass Phalaris arundinacea reed canary grass Eragrostis cilianensis stinkgrass