ANATOMY AND MORPHOLOGY OF THE

POACEAE [GRAMINEAE] GRASS FAMILY

Slide 2 A quick family description as found with all family files on this site.
Slide 3 A brief comparison of Graminoids (grasses, sedges and rushes)
Slide 4 to 43 - Anatomy and Morphology of Grasses

SOME HINTS, HELPS, AND THOUGHTS ON WORKING WITH GRASSES

1. -One must examine the roots (and rhizomes, etc. if present) of a grass to determine if it is an annual or a
perennial. I carry a long Philips-head screwdriver for digging purposes (easier to carry than a digging tool)
2. -In most cases one must examine a grass when in flower (Anthesis) to best be able to key it out (occasionally the
fruiting stage is important)
3. -Although some can ID grasses in the field using a hand lens, an amateur (and sometimes the expert) must see the
spikelet structure, etc. under a binocular microscope (often with a little dissecting). Once thoroughly familiar with a
species most grasses can usually then be ID in the field.
4. -To keep samples fresh for 1 or 2 weeks or more I use Ziplock bags (usually gallon size) and in the field I mist them
using a small plastic pump bottle with water and then keep the bags in the refrigerator at home.
5. -The point-and-shoot cameras of today makes recording a plant easy. I try to take pictures (weather I use them or not)
if it may be a new species (for my database) of all features using the close-up mode (often the light is better in the
field than at home). At home I can also take pictures (with patience) through the eyepiece of my old binocular
microscope (this works amazing well without having an expensive lab instrument).
6. -At first it was very difficult using the keys (and occasionally it still is) but with experience in observing and describing
the array of grass structures, it gets easier especially as one can often determine the Genus and work the keys from
there.

POACEAE [GRAMINEAE] GRASS FAMILY

Plant: annuals or perennials
Stem: jointed stem is termed a culm internodial stem most often hollow (may have
pith) but always solid at node, mostly round (some exceptions), some with stolons
(creeping stem) or rhizomes (underground stem)
Root: usually fibrous, often very abundant and dense
Leaves: mostly linear, sessile, alternate, with parallel veins, in 2 ranks (vertical rows),
leaf sheath often open to the node (may be overlapping) or partially fused (closed)
Flowers: small in 2 rows forming a spikelet (1 to several flowers), may be 1 to many
spikelets with pedicels or sessile to stem; each flower within a spikelet is between an
outer lemma (bract, with a midrib) and an inner palea (bract, 2-nerved) these 3 parts
together make the floret the 2 bottom bracts of the spikelet do not have flowers and
are termed glumes (may be reduced or absent), the rachilla is the axis that hold the
florets; sepals and petals absent; 1-6 but often 3 stamens; 1 pistil, 1-3 but usually 2
styles, ovary superior, 1 ovule
Fruit: seed-like grain (seed usually fused to the pericarp (ovary wall) or not)
Other: very large and important family; Monocotyledons Group
Genera: 600+ genera; locally many genera
WARNING family descriptions are only a laymans guide and should not be used as definitive

GRASSES, SEDGES AND RUSHES (sometimes termed Graminoids)

[V. Max Brown]

Grasses (Gramineae or Poaceae), Sedges (Cyperaceae) and Rushes (Juncaceae) are monocotyledon
plants that at first glance may seem to look alike but are mostly separated fairly easily. However, exceptions
do occur for many characteristics below so beware!!! All three families have leaves with parallel veins, one
seed leaf and the vascular bundles are scattered within the pith of the stem.
Grasses Plants annual or perennial, herbaceous to rarely woody, caespitose (forming dense tufts) or not, perennials usually
with rhizomes and sometimes stolons; Roots fibrous; Leaves basal and/or cauline, 2-ranked (vertical rows), alternate, mostly
flat (may curl or be folded), leaf sheaths usually open or split (but often overlapping), sometimes sheaths are partially to mostly
closed or fused, Ligules often present, auricles sometimes present; Stems (culms) are usually round (may be oval or
somewhat flattened), nodes solid and usually swollen, internodes hollow or solid; Inflorescence of small Florets (flowers)
within Spikelets (subtended by 2 empty bracts termed Glumes) forming Spikes, Racemes, or Panicles or some combination
thereof; Florets (flowers) usually enclosed in 2 bracts (Lemma and Palea), petals and sepals absent, with usually 3 stamens,
1 pistil; Fruit usually a Caryopse (Grain), seed fused to pericarp.
Sedges Plants annual or mostly perennial, herbaceous, caespitose (forming dense tufts) or not, perennials usually with
rhizomes and sometimes with stolons; Roots fibrous; Leaves basal and/or cauline, alternate, mostly 3-ranked (vertical rows)
but not always, blades sometimes flat or V- or M-shaped, rarely round or reduced, midvein usually large, leaf sheaths usually
closed or fused, blades may be absent in basal leaves, ligules often present but may be reduced; Stems (culms) most often
triangular but some are round (terete), sometimes flattened, rarely 4- or 5-angled, nodes lacking, interior usually solid (Pith);
Inflorescence of one to many terminal and/or axillary (rarely basal) spikelets in panicles, umbels, racemes, spikes or headlike clusters (capitulum) which may be subtended by bracts; Flowers mostly bisexual (some unisexual), perianth absent
(naked) or reduced to bristles or scales, 1 to 3 stamens, 1 pistil (enclosed in a sac-like structure (perigynium) in the genus
Carex), ovary superior; Fruit a biconvex or trigonous achene,1 seed free of pericarp.

Rushes Plants mostly perennial, occasionally annual, herbaceous, caespitose (forming dense tufts) or not, mostly with
rhizomes; Roots fibrous; Leaves 3-ranked (vertical rows) but rarely 2-ranked, mostly basal, often round (terete) but
sometimes grass-like (flat) or reduced, leaf sheaths usually open or split, some with ligules and/or auricles; Stems (culms) are
usually round (terete), sometimes flattened, occasionally 4- or 5-angled, nodes lacking, interior solid; Inflorescence of
spikelets in compound panicles and racemes of cymes, umbels or spikes of head-like clusters to single terminal spikelets,
often subtended by bracts; Flowers perfect, mostly bisexual, occasionally unisexual, perianth present usually as greenish to
brownish tepals (3+3), 3 or 6 stamens, compound superior ovary, 3 stigmas; Fruit a capsule with 3 to many seeds.
**A common saying (but user beware) sedges have edges, rushes are round, grasses are hollow right up from the ground
** A variation Sedges have edges and rushes are round, grasses have joints when the cops are not around

GRASS ANATOMY Typical Grass

[V. Max Brown]

Inflorescence the
flowering and seed
forming (reproduction)
part of a grass plant

Culm - (stem of
grass below the
inflorescence)
Leaf - leaves 2ranked (occur on
opposite sides of
culm), and arranged
alternately on culm

Roots Fibrous
roots

Although many terms will be

defined, knowledge of some basic
botanical terminology is expected

GRASS ANATOMY Culms, Nodes and Internodes

[V. Max Brown]

Culm the jointed flowering stem of

grasses (usually Herbaceous but
sometimes (rarely) Woody)
Nodes (Joints) are solid in
cross-section and often are
somewhat swollen due to
growth of the leaf base.
Internodes (between nodes) are
usually hollow (sometimes oval or
flattened, or with pith) and may be
glabrous or hairy, rough, ribbed,
etc.
Leaf Blade
Sheath of Leaf Blade
wraps around culm in
many grasses

Culm, leaf blade and

sheath with dense
spreading hairs

Culm with strongly

(upward) appressed
hairs

Culms may be Erect (standing) or Decumbent (prostrate or lying down)

or almost anything in between. Culms may branch either at the Basal
Nodes only or at Base, Middle and Upper Nodes. If branches are
mostly basal the grass often forms Tufts. Lower portion of culms may be
differently colored or shaped than the upper culm in some species.
Caespitose (Cespitose) growing in dense tufts (dense clusters)
Pluricespitose many dense tufts
Innovations short non-blooming basal stems of some grasses

Culms flattened near

base in a few grasses

GRASS ANATOMY Culms, Nodes and Internodes

[V. Max Brown]

Internode of culm hollow;

vascular bundles in walls; this is
an example of a somewhat
flattened (compressed) culm

Internode
flattened, not
hollow but
with spongy
pith

Node solid,
internode
hollow

Node swollen,
Node swollen,
reddish, internodes glabrous (smooth),
mostly smooth
internodes of culm
ribbed

Keeled - Culm and leaf

sheath flattened to a ridge
like a Keel of a boat

Glands
(rare)

Node slightly swollen,

glabrous (smooth); node
and internode area below
glaucous (with white
waxy coating usually
easily rubbed off)

Node swollen, with

appressed
(upward) hairs

Node swollen, with

appressed (downward)
hairs, leaf sheath above
with prickles

Nodes long
retrorsely bearded
(tufts of long hair)

GRASS ANATOMY Leaves

[V. Max Brown]

Foliage Leaf the numerous large leaves of grasses are alternate and 2-ranked on the Culm, and originate at the
node.
Foliage Leaf Blade usually narrow and long (linear to lanceolate) but sometimes ovate, triangular or other
shapes. They have Parallel veins (with rare exceptions) and the tip or apex of the blade is often fairly sharp-pointed.

Linear to Lanceolate

Somewhat ovate
to lanceolate leaf

There is no true Petiole in Grasses

but some have a constriction
(Pseudo-Petiole) and is termed
Pseudopetiolate.

Crown the base of a herbaceous (non-woody) perennial plant that tends

to persist from year to year; shoots from the Crown can cause thick tufts to
form.

GRASS ANATOMY Leaves

[V. Max Brown]

Grass leaves may be Basal and/or Cauline (stem). In some species the cauline leaves die back
with winter and a winter basal rosette of leaves is present.

Mostly cauline leaves, a

few basal leaves

Both cauline and basal

leaves common

Cauline leaves with a basal

rosette of leaves left over from
previous winter (these rosettes
when present may or may not be
similar to the cauline leaves)

GRASS ANATOMY Leaves

[V. Max Brown]

A Foliage Leaf is attached to the Culm (stem) by a Sheath that is ultimately attached at its base,
the Node (leaves originate at nodes). In many grasses, the Sheath is Open (split or unfused) and
often partially overlaps around the Culm however, in some the sheath is fused (closed) partially
to almost totally.

Node

Culm

Leaf Blade

Leaf Blade Base

Leaf
Sheath
grows from
node

Culm
Leaf Sheath

Leaf Sheath
is here pulled
away from
culm

Node

Culm

GRASS ANATOMY Leaves

[V. Max Brown]

Midrib the larger central vein or rib of the leaf, usually there are several smaller veins or ribs as well although grasses
have Parallel veins there may be smaller cross-veins present and occasionally veins may diverge. Leaf blades may be
glabrous (smooth), hairy, etc. as well.
In the photos below, AD (Adaxial) upper surface facing culm and AB (Abaxial) lower surface facing away from the culm

Many fine ribs, no

prominent midrib (AD)

Some long hairs (AB)

4 prominent ribs on
each side of larger
midrib (AD)

Abundant Long hairs

(AD) and on sheath

Very prominent white midrib

(AD)

Dense long hairs on base

of leaf blade (AD)

Prominent rib of midrib

(AB)

Midrib (vein) off-center

in leaf blade (AD)

Fine prickles on
leaf edge

GRASS ANATOMY Leaf Sheath open and closed

[V. Max Brown]

The Leaf Sheath may be Open (unfused) all the way down to the Node or it may be fused at any point
from the Node upward toward the base of the leaf blade.

Closed to above
middle
Leaf Sheath is open all the way to the node
however the sides of the leaf sheath will often
overlap as in some of the above examples
(open is a common condition)

Sheath closed near

top of sheath, some
call this the V
necked sweater
condition

Leaf sheath Closed Sheath may be fused from the node up only a
little ways up the sheath to far up near the base of the leaf blade (almost
completely fused). It is often important in keying out some grasses to
know not only if it is closed but if it is closed below the middle (a little)
or above the middle (a lot). etc.

GRASS ANATOMY Branches

[V. Max Brown]

Branches (if present) from the Culm originate

at the node, usually above the base of the leaf.
There are 3 possibilities as to how the branch
emerges in relationship to the Culm and the
Leaf Sheath.

1. Intravaginal branch emerges between

the leaf sheath and Culm at the base of the leaf
blade (if near plant base Tillers are developed).
A Tiller is a basal or underground Shoot (young
stem or branch) that is growing fairly upright
(erect) at the base.

Intravaginal

Branch
Leaf
Blade
Culm

2. Extravaginal - branch emerges by piercing

the basal portion of the leaf sheath (if near base
or Crown, Rhizomes or Stolons may develop).
3. Infravaginal branch is within the leaf
sheath? Condition is rare
Basal Leaves branches may be Intravaginal
or Extravaginal
Upper Node Leaves branches may be
Intravaginal, Extravaginal, or Infravaginal
Prophyllum or Prophyll (Bracteole)
modified first leaf of a shoot or branch at a
node of the Culm The Prophyllum lies
between the culm and the branch on the main
branch side and braces the branch, it is often
H shaped.

Branch
Leaf
Sheath

GRASS ANATOMY The Ligule

[V. Max Brown]

Ligule - a small membrane or sometimes just a line or fringe of hairs, or Ciliate (membranous with cilia of hairs) on the
sheath at the junction of the blade and the sheath in many but not all grasses (may be absent). The Ligule (sometimes
referred to as the rain-guard protecting the sheath) is on the Adaxial side of sheath (toward axis or Culm in this case)
Abaxial would be the opposite orientation.
Culm or stem has been removed in most photos below

Ligule absent (there is no

extra tissue here)

Ligule mostly of
hairs

Ligule membranous,
margin even

Ligule membranous, margin

with pointed or acute tip

Ligule membranous, erose

(looking torn or gnawed, uneven
or toothed)

Ligule membranous and ciliate (with hairs on edge of membrane)

A rare Ligule of
bristles

GRASS ANATOMY Auricle and Collar

[V. Max Brown]

Auricles (Auriculate) small ear-like projections at the junction of the Leaf Sheath and the base of the Leaf of some grasses;
Collar the area found at the junction of leaf sheath and base of leaf blade on Abaxial side (opposite side to Ligule and culm).

Leaf Blade

Auricle
Leaf
Sheath

Auricle
Auricle
Auricle
Leaf Blade
Leaf
Sheath

Collar

Collar the area on the

opposite side to the ligule at the
junction of the leaf sheath and
leaf blade the collar may be
hairy or glabrous, etc.

Collar rounded

Auricle

GRASS ANATOMY Fibrous Roots

[V. Max Brown]

Fibrous Roots besides the short-lived primary root that grows from the seed, all grasses have fibrous roots
(roots that are small in diameter and usually in large numbers) - in addition, some grasses have underground
stems (Rhizomes) and above ground runners (Stolons) (next slide).

It is important in most ID keys to know if a grass is

an Annual or a Perennial.
An Annual is a plant that germinates from seed,
grows and flowers, sets seed, and dies all within one
year. Next year plant again develops from seed.
A Perennial is a plant that lives multiple years.
To tell the difference is not always easy if the plant
has a soft base (plant with shallow fibrous roots that
pulls up easy) it is probably an Annual.
If it has a hard base (difficult to pull up, often having
deeper fibrous roots and Rhizomes) and sometimes
still has old growth from previous year(s) it is probably
a Perennial.

* One should always examine the roots of a

grass when making an ID an old
screwdriver, large knife or other type of
digging tool is often required.

GRASS ANATOMY Stems other than Culms

Stolon (runner) a stem that creeps along
the ground, often rooting at the nodes

[V. Max Brown]

Rhizome an underground stem, sometimes with Scale Leaves

(reduced leaves found near plant base or on rhizomes)

Scaly Rhizomes

Roots at node of Stolon

Tiller a Tiller is a basal or underground Shoot

(young stem or branch) that is growing fairly
upright (erect) at the base

Large, non-scaly Rhizomes

Large, short Rhizomes

GRASS ANATOMY Inflorescence (Flowering Portion of Plant)

[V. Max Brown]

Rachis the central stalk within an Inflorescence; Rachilla a secondary internode or branch within the inflorescence
Spikelet the basic unit (ultimate or smallest flower cluster) of a grass. An inflorescence is made up spikelets with flowers.
Pedicel the stalk which holds a Spikelet unless it is attached directly to a branch or the Rachis and is then said to be
Sessile (no stalk).
Peduncle the stalk just below the Inflorescence

Inflorescence
Unbalanced
Spike - to
one side

Spikelet
Rachis

Spikelet

Pedicel

Rachis
Culm

No Pedicels above
because spikelets
are Sessile on the
Rachis also, it is
a Balanced Spike

GRASS ANATOMY Inflorescence Types

[V. Max Brown]

The Inflorescence of a grass may be a:

1. -Spike - a simple (single axis) spike with spikelets sessile to Rachis. Spikelets may be all on one side of
axis, symmetrical along axis (alternating on both sides of Rachis a Balanced Spike or surrounding the
Rachis) or other configurations (be careful in grasses, what may seem to be a Spike at first glance is often a
Raceme or even a contracted panicle!)
2. -Raceme - a spike with the spikelets on pedicels of varying length, flowers lower on the Inflorescence
maturing first
3. -Panicle - a branched Raceme (very common inflorescence type in grasses) panicles can vary from quite
open to very dense or compact. Often even if the branch is a spike it is still termed a Panicle in other
words, if the Inflorescence is branched it will probably be termed a panicle with qualifiers.
4. -and other compound and complex arrangements of spikelets (such as RAMES discussed later in spikelet
examples). Often it seems easier sometimes to describe some of these as spike-like or spicate or racemelike or racemose branches of panicles, etc.

GRASS ANATOMY Inflorescence - Spikes

[V. Max Brown]

1. -Spike - a simple (single axis) spike with spikelets sessile to Rachis. Spikelets may be all on one side of axis, symmetrical
along axis (Balanced) or other configurations (be careful in grasses, what may seem to be a Spike at first glance is often a
Raceme!). Below are 3 examples of Balanced Spikes and 1 one-sided spike.
Thinopyrum ponticum
(Podp.) Z.-W. Liu & R.-C.
Lolium perenne L. (Introduced)
Wang (Tall Wheatgrass)
(Perennial [English] Ryegrass)
(Introduced)

Bouteloua hirsute Lag. (hairy Grama)

1-sided spike
Hordeum pusillum
Nutt. (Little Barley)

Although this is termed a spike

in this species, some might say
it is a spike-like raceme.
Although difficult to see here the
spikelets are attached on
opposite sides of the
inflorescence
Spikelets attached edgewise to the culm culm
is indented here

Spikelets attached by
the flat side to the culm

GRASS ANATOMY Inflorescence - Racemes

[V. Max Brown]

2. -Raceme - a spike with the spikelets on pedicels of varying length, flowers lower on the Inflorescence maturing first;
Vulpia myuros (L.) C.C. Gmel.
(Rattail [Annual; Foxtail] Fescue)

Danthonia spicata (L.) P. Beauv. Ex

Roem. & Schult. (Poverty Oatgrass)

Spikelet

Pedicel

GRASS ANATOMY Inflorescence - Panicles

[V. Max Brown]

Phleum pratense L.
Introduced (Timothy)

3. -Panicle - a branched Raceme (very common inflorescence type in grasses)

and particularly in grasses a spike-like Raceme (see timothy, to right).

Bromus tectorum L.
(Introduced) (Downy
Brome[Cheatgrass])

Dense Panicle

Panicum virgatum L.
(Switchgrass)

Open Panicle

Dichanthelium boscii (Poir.)

(Bosc's Panicgrass)

Very Open Panicle

Contracted Panicle Very dense or compact

GRASS ANATOMY Inflorescence - Pulvini

[V. Max Brown]

Pulvini are swellings (usually at the base of a petiole) but here at the base of a branch
from the Rachis in a grass Inflorescence. Sometimes the characteristics of these
Pulvini may aid in ID of some species.

Pulvini with
dense hairs
covering the
swellings
Pulvini with
very long hairs

Pulvini with
hairs only on top
of swelling

GRASS ANATOMY Spikelet Anatomy and Morphology

[V. Max Brown]

Most of the botanical terms need to describe Spikelets is given below and on the next slide. These
terms and others will be presented with illustrations and descriptions of a few individual species to
follow
Each Spikelet often contains 2 basal Glumes (empty Bracts, 1 on each side of axis) with 1 to 60+ Florets (flowers) above the
Glumes which are attached to the Rachis (central axis) or Rachilla (a branch, smaller order or little Rachis) in a compound
Inflorescence. The Glumes and Florets are attached Distichously (Distichous) (2-ranked in 2 vertical rows on opposite
sides of axis). A Glume typically has an odd number of veins and may have an Awn(s) - a stiff bristle at tip. A Glume may vary
in shape and may look like a Awn itself (Subulate). Veins and awns may be important in ID. In some species 1 or both
glumes may be absent.
A Floret (flower) consists usually of a Lemma (lower outer bract), a Palea (upper inner Bract) with the reproductive apparatus
(stamens and pistils between them). Lodicules (0, 2 to 3) are scales or swellings at the base of the ovary (may represent
what's left of the Perianth of flower) that at flowering swells and forces the Lemma outward to expose the reproductive parts of
the flower to the wind. No Sepals or Petals (Perianth) are present and so the flower is said to be naked.
A Lemma (lower and outer bract of a floret) has veins (usually an odd number with a more prominent midrib and lesser veins
parallel to it but they do tend to converge toward the tip). Often, the Midrib forms a Keel (Lemma is folded or rounded over the
Midrib). The Lemma may have 1 (less often 2 or 3) Awns (bristles) usually arising from an extension of the midrib (and less
often from other veins) awns (when present) may originate from near the base, middle, or near tip of Lemma. It also has a
hard and thick base termed a Callus (may be well developed or not and hairy or not).
The Palea, (upper and inner bract of the floret) is occasionally absent or is very small in some grasses, however it is present in
most grasses (sometimes transparent) and it has 2 major veins, usually 1 each along a margin (sometimes ciliate) and may or
may not have other minor veins.
Each Floret (flower) may be Bisexual (Perfect, male and female with both stamens and pistils); Or Staminate only (male,
having Stamens only and will not bear fruit); Or Pistillate only (female, having pistils but no stamens); Or Sterile (having
neither staminate or pistillate structures Or having a reduced structure); Or a spikelet with multiple florets may have mixed
florets of above. Also, a few grasses are Monoecious and a few Dioecious.

GRASS ANATOMY Spikelet Anatomy and Morphology

[V. Max Brown]

Botanical Terms continued

There are usually 3 Stamens (sometimes 1 or 2 or 6+), usually alike but sometimes 1 or more are longer than the others.
Anthers attached mostly to filaments in the middle (Versatile).
The Ovary has 1 locule (cavity), 1-4 (usually 2-3) styles, styles may be branched, Stigma usually Plumose (feathery).
Caryopis the grass fruit is 1-seeded, dry, and indehiscent (not breaking open along predefined lines or pores) kind of
achene or Grain. The seed is fused to the fruit or ovary wall (Pericarp). An Endosperm is present and is attached to the
embryo and is used as food.
At maturity the Spikelet disarticulates (breaks) to allow the spread of the seeds. The Spikelet may break off below the
Glumes leaving an empty Pedicel or it may break above the Glumes (leaving them on the plant). In the latter case, the
Spikelet usually also breaks up between the Florets. This may be important for ID in some keys.
Some Spikelets are round in cross-section but others may be compressed either laterally (from the sides of the glumes,
further folding them with more pronounced keels) or dorsally (from the backs of the Glumes, they are then more opened)

There is large diversity in the anatomy of Grasses. The plants may be erect to decumbent, small to large, leaves may vary
from small to large and in various shapes, the sheath is usually open but may be partially to wholly closed, the Glumes of a
Spikelet may be severely reduced to almost nothing or so large they cover the Florets, in some the upper Florets are fertile
and the lower Sterile, and some Spikelets may be male only or female only and in some grasses a plant may be male only or
female only. A few grasses are Cleistogamous (self pollinating). In addition many parts of grasses may or may not have
bristles, hairs, awns, keels, etc. At first grasses may seem to all look alike but they do vary widely as one should expect
from the 4th largest family of flowering plants.

GRASS ANATOMY Spikelet Anatomy and Morphology - General Look

[V. Max Brown]

Spikelet

Spikelet

Awns from
Lemmas of
Florets

Upper Florets
yet to open
Only the Lemma of
each floret is in view
(Palea is hidden).
The Spikelet is very
Dorsally flattened,
giving the Lemmas
well developed keels

10
Florets

Stamens from
the opening of
a few lower
florets

Upper Glume

Stigmas of
lower open
florets

Upper Glume

Lower Glume

Lower Glume
Pedicel

The photos above illustrate rather simple spikelets with glumes and multiple fertile florets. There
are many exceptions to this as you will see in the following slides. It is best if one quickly
memorizes the terminology of grass anatomy before continuing.

GRASSES SubFamilies, Tribes, Genera and Species

[V. Max Brown]

The Flora of North America, Volumes 24 and 25, lists 10 Subfamilies, 25 Tribes,
236 Genera and 1,373 Species in North America (North of Mexico) for
comparison sake, The Flora of Missouri, Volume 1, lists 89 Genera and 276+
Species - 700+ Genera and 11,000+ Species estimated world-wide.
Tribes are important in ID grasses and most keys use Tribes. However, not all
sources agree on the number and names of grass Tribes present. On this website
I have used the Tribes as listed in the Flora of North America.

The following slides will try to illustrate some of the diversity of Spikelet Morphology by
looking at examples in several Tribes. Measurements are often critical in ID but they are
mostly not given in these examples (see individual species in database).

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Panicum virgatum L. (Switchgrass) Paniceae Tribe

1.
Pic 1. -a single Spikelet with 2 Glumes, it also has 2
Florets (see Pic 2. below)
Florets

Lower Glume

Pic 2. -in this view the lower glume is missing

(removed) and the spikelet is spread apart showing the
lower Staminate floret (male only, anther present) and
the upper Perfect (fertile) Floret. The upper floret is
glabrous and shiny and its Lemma and Palea cannot be
distinguished in this photo. Note that the first Lemma
(here the lower floret lemma) is opposite to the upper
glume which is opposite to the lower glume.

Upper glume

Lower Staminate Floret

2.

Sometimes the number of veins in the glumes and

lemmas may occasionally be important in ID

Upper Fertile
Floret
Anther
Lower
Floret Palea

Lower
Floret
Lemma

Upper Glume, lower Glume

missing in this photo

This densely hairy (Pilose)

triangle of long hairs at base of
leaf sheath (just above Ligule)
can often be used for a quick
ID of this species if other
properties such as height,
inflorescence type, etc. fit.
Some authors say it is
occasionally absent?

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Setaria parviflora (Poir.) Kergulen (Marsh Bristle

grass [Knotroot Foxtail]) - Paniceae Tribe

1.
Note the transverse
wrinkles (rugose
texture) on the upper
Lemma

Pic 1. The Spikelet is ovoid in shape and though not seen

here there are 2 florets, a lower, usually staminate floret and
an upper perfect floret dissection under a scope is needed
to see this.

Lower
Glume
(longer)

Upper
Glume
(smaller)

The spikelets are in dense clusters on very short branches

so, the inflorescence in this case is is a dense, spike-like
Panicle (again, difficult to see in the field).
Pic 2. The anthers and stigma protrude from the spikelet
without clearly showing the 2 florets only dissection will
show this.

Spikelet subtended
by 4 to 12, usually
yellowish bristles,
antrorsely (forward
directed) barbed

2.

Anthers
Inflorescence
axis with
short hairs

Feathery stigma

In addition the fact that this species is a perennial with

short rhizomes and the size of spikelet and number of
bristles was important in using a key for ID

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Pic 1. This species is an example of a grass with Fascicles (Fascicle is a

general term meaning a tight cluster) a hard bur-like structure in grasses
derived from branches of the panicle. In this species It has very sharp, hard
flattened bristles, is densely hairy, and surrounds 1 or rarely 2 spikelets with 2
florets 1 sterile and 1 perfect.

Cenchrus tribuloides L. (Dune

Sandbur) - Paniceae Tribe

Pic 2. The dried Fascicles can make for painful walking on the beach. This
feature is always early in any grass ID key.

1.

2.

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Dichanthelium boscii (Poir.) (Bosc's Panicgrass) - Paniceae Tribe

Spikelet with lower staminate floret and upper fertile floret, spikelet length in this
case 4.5 mm (can vary from 3+ to 5+ mm)

1.

Upper Glume

Pic 1. (top left 3 pictures) The spikelet has a

small lower glume and a large upper glume.
There is a lower staminate floret and an upper
perfect or bisexual floret. The spikelet is
Puberulent (short hairy) and usually, like the
Genus Panicum, is fairy rigid or hard.
Pic 2. Lower staminate floret with anther
shown, removed from spikelet. Upper bisexual
floret removed as well, note the tiny tuft of hair
and smooth Lemma.

Lower Lemma

Pic 3. inflorescence a Panicle

Upper Glume,
slightly shorter
than spikelet
Lower Glume

3.

2.
Tiny tuft of
hair often
Lower
staminate floret
with
transparent
palea

Upper fertile
floret
removed
from spikelet

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Phleum pratense L. Introduced (Timothy) Poeae Tribe

Pic 1. the Spikelet is small, with 2 glumes of about
equal length that are much longer than the 1 floret
hidden inside. The sides of the glumes are Puberulent
(finely hairy) and the keels are long ciliate. The glumes
have short awns at their tips.

1 Spikelet with 1 fertile floret inside

1.

glumes

Pic 2. The single Perfect Floret (here removed from

the glumes) is immature but the Lemma and Palea are
visible and at least the anthers are recognizable.
Pic 3. A close-up view of the spikelets show that this
is actually a Panicle (Inflorescence) the florets are
really on very small branches that are fused (adnate)
to the rachis.

Long Hairy on
keel of glumes

2.

The inflorescence, spikelet shape and tall ligule is

usually enough to guess that this is Timothy although
there are other species in this genus that are somewhat
similar

1 Floret
anthers

3.
Palea

Lemma

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Poa bulbosa L. (Introduced) - Bulbous Bluegrass Poeae Tribe

A.
Subspecies bulbosa
with normal fertile florets
(note the bulb-like culm
bases)

B.
Subspecies Vivipara
with vegetative
bulblets replacing the
normal fertile florets,
these bulblets will drop
off and prorogate the
species

Vegetative
Bulblet

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Diarrhena obovata (Gleason) Brandenburg (Obovate [American] Beakgrain) Diarrheneae (Tribe)

1.

Upper
sterile
floret

Pic 1. Full spikelet in flower with 2

glumes( lower one small) and 4 florets (3
perfect florets and 1 upper sterile floret).
The Lemmas have short awns. The
Paleas are tucked into the Lemmas and
are not shown
Pic 2. Two lower mature fertile florets
shown (removed), note how the
Caryopis (fruit) is pushing the Lemma
and Palea apart.

2.
Lemma
of Floret

Upper glume

Awn of
Lemma
Caryopis

3.

Pic 3. one very mature floret pulled

from spikelet. The Caryopis has a very
abruptly narrowed neck or beak with a
notched tip another very similar
species has a gradually tapering neck.
In this case a mature fruit is necessary
for easy ID

Upper glume
Lower Glume

Lemma
Palea

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Leersia virginica Willd. (White Grass [ Cutgrass]

- Oryzeae Tribe (rice is in this tribe)

1.

Spikelet (opposite views)

3 mm

Pic 1. - This is an example in which the usual glumes are

reduced to a cup-like structure. There is 1 floret. The Lemma
curves around and the palea is somewhat inset. The veins of
the Lemma are usually ciliate.
Palea (3
nerved),
often
with cilia
on edges

Glumes
reduced to a
cup-like
structure at
base of spikelet

Lemma (5
nerved,
ciliate or not)

Leaf blades and sheath often rough with small bristles.

Pic 2. the spikelets are loosely overlapping but not tightly so.
The Inflorescence is a panicle. The size and shape of the
Spikelet, lack of glumes, etc. is very important in using the ID
keys.

2.

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Aristida longespica Poir. (Slimspike Three Awn) Aristideae (Tribe)

1.
Small awns
on Glumes
Upper Glume

Lemma
Lower Glume

Pic 1. and Pic 2. There is 1 floret per spikelet, the glumes

(also awned) are about as long as the body of the Lemma.
The Lemma has 3 awns that bend on drying. The Palea is
much smaller than the Lemma. A sharp Callus (extension of
the Lemma) is present.
Pic 3. an immature spikelet (before drying and bending of
awns.
The length of the Lemma awns, and whether they are twisted
or flattened, etc. are important in ID keys.

2.

Awns bent on drying

but not twisted
(spiraled) or overly
flattened at base
(other species)

Callus

3.

Lateral awns of Lemma are

shorter than central awn In this example the central
awn is 22 mm, both lateral
awns are about 14 mm,

Mature
spikelet, post
flowering

Immature
spikelet from
lower part of
inflorescence,
awns not
dried and
bent yet

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Sorghum halepense (L.) Pers. (Introduced)

(Johnson Grass) - Andropogoneae Tribe

Spikelet Pairs and the RAME Structure in the

Andropogoneae Tribe

Another pair

Grasses in the Andropogoneae Tribe usually have spikelet pairs (a

few have 3 spikelets) and each spikelet has 2 florets these spikelet
pairs make up a unit or basic structure for description. The glumes
completely enclose the florets.
In most cases one spikelet is sessile (usually holding an upper fertile
floret, and lower staminate or sterile floret) and the second spikelet is
pedicellate (on a pedicel) and its florets are usually staminate, but often
sterile, reduced or even absent (only the pedicel is left in some cases).

pedicellate
(staminate)
spikelet of
2st pair
sessile
spikelet (with
perfect floret)
of 2st pair

pedicellate
(staminate)
spikelet of
1st pair

Pedicel of
pedicellate
spikelet (2nd
pair)

Pedicel of
pedicellate
(staminate)
spikelet (1st
pair)

Branch or
internode to
next 2-spikelet
pair

sessile spikelet
(with perfect floret)
of 1st pair

A unit of structure is then both the sessile and pedicellate spikelets plus
the branch or internode that goes up to the next sessile spikelet of the
next pair of spikelets (disarticulation often taking place here).
A RAME (term of structure) is an inflorescence or part of an
inflorescence composed of the basic units disarticulating as above. I
have read descriptions in which the term RAME seems to be used for
the basic 2-spikelet structure.
If the sessile and pedicellate spikelets are sexually different they are
called heterogamous; if sexually alike they are called homogamous.
Most grasses in the Andropogoneae are heterogamous. If the sessile
and pedicellate spikelets are morphologically different they are called
heteromorphic; if morphologically alike they are called homomorphic.
Most grasses in the Andropogoneae are heteromorphic.

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

The 4 examples below illustrate some

of the features of the 2-spikelet units in
the Andropogoneae Tribe although
there is still a lot of diversity not
illustrated. Note the hairiness of many
of the samples. A RAME inflorescence
to the right. The next slide illustrates a
monoecious example from the
Andropogoneae.

Pic 1. - Saccharum alopecuroides (L.) Nutt.

(Silver Plume Grass)
Pic 2. - Schizachyrium scoparium (Michx.)
Nash var. scoparium (Little Bluestem)
Pic 3. - Bothriochloa laguroides (DC.)
Herter (Silver Beardgrass)
Pic 4. - Andropogon virginicus L. var.
virginicus (Broomsedge Bluestem)

2.
Pedicellate
spikelet

Upper sterile
or staminate
spikelet

1.

Upper floret
lemma of
sessile spikelet
with awn
somewhat
spiraled and
bent

A RAME Inflorescence

4.

3.

Pecidellate sterile
spikelet absent
or much reduced
Sessile
Spikelet

Pedicel about as
long or less than
sessile spikelet

Pedicellate
Spkelet,
reduced and
sterile, shorter
than sessile
spikelet

Sessile
Spikelet

Pedicel of sterile spikelet,

usually about twice as
long as sessile spikelet

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Tripsacum dactyloides (L.) L. (Eastern Gama Grass) Andropogoneae Tribe

Pic 1. Eastern Gama Grass is

Monoecious with Staminate Spikelets
above and Pistillate Spikelets below
(2 florets each) on each RAME
(typically 2 or 3 RAMES are present).
The basic 2-spikelet units are a little
different in monoecious grasses as
compared to perfect examples.

2.
1.
Paired Staminate
Spikelets, sessile
or with very short
pedicels, stamen
anthers shown
here

Pistillate Spikelets,
Lower Glume
covers 2 florets, 1
sterile (lower) and 1
fertile (upper)
pistillate florets,
Feathery styles of
pistillate floret

Pic 2. immature staminate spikelets

Pic 3. immature pistillate spikelets;,
lower inset shows styles

3.

inflorescence

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Elymus villosus Muhl. ex Willd. (Hairy [Downy] Wild Rye) Triticeae Tribe

1.

Pic 1. There are 2 spikelets at each node with 1 or 2 (rarely 3)

florets per spikelet (the lower floret of each spikelet is usually the
fertile or functional one). In this example there are 4 glumes (2
per spikelet and awned), all of which are densely hairy (villous).
The Lemmas also have awns and are villous. The florets are
not dissected or shown below.
Note in this species that the lower part of the glumes are straight
sided and not bulged may be important in ID

Glumes

spikelets

Paired spikelets rotated to look at

(abaxial) side (away from culm)

Nodes with
paired
spikelets
removed

A Lower
Glume

Paired spikelets rotated to look

at adaxial side (toward culm)

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Hordeum pusillum Nutt. (Little Barley) Triticeae Tribe

3 Spikelets at node, 1 central and 2 lateral

Adaxial view of triple spikelets
at node (toward rachis)

1.

Abaxial view of
tripe spikelets at
node (away from
rachis)

Glumes

2 smaller lateral
spikelets

1 Larger central
spikelet

Pic 1. There are 3 spikelets at each node (sessile

to pedicellate) and 1 floret per spikelet in this grass
from the Triticeae Tribe. The larger central spikelet
is usually sessile with a bisexual (fertile) floret and
the 2 lateral spikelets usually pedicellate and have
either sterile or reduced florets. Glumes and
Lemmas are usually awned. Note that the glumes
become flattened toward their base.

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Bromus hordeaceus L. (Introduced)

(Soft Brome [Chess] [Lopgrass])
Bromeae Tribe

Pic 1. This species has a spikelet with 2 glumes and 5 to 10

perfect florets, and the Lemmas are awned. Both glumes and
Lemmas finely hairy.
Pic 2. the inflorescence is fairly compact, spikelet stalks
shorter than spikelets.
Pics 3. note the awn from the Lemma midrib and the apical
teeth

Apical teeth
of Lemma

3.

1 Floret

1.

Awn is an
extension of the
Lemma midrib

Florets

Caryopis
Palea (pulled out
some) nearly
transparent with
ciliate margins,
shorter than
Lemma

2.

Lemma
Glumes
Transparent Palea

GRASS ANATOMY Spikelet Morphology - Examples

[V. Max Brown]

Melica nitens (Scribn.) Nutt. ex Piper (Three-Flowered Melic Grass) - Meliceae Tribe

Club-shaped Rudiment
in sterile floret

Upper Sterile Floret

2.

1.

Pic 1. In this species there are 2 to 3 (rarely 4)

florets above the glumes. They are all perfect
(bisexual) except the top one which is sterile. In the
Genus Melica this reduced structure is termed a
Rudiment. In the Genus Melica as a whole there
may be up to 7 bisexual florets and up to 4 sterile
florets (with rudiments) in various species.
Pic 2. the club-shaped Rudiment.
Pic 3. the inflorescence is a Panicle

Fertile florets not

at same level in
spikelet

Note Rachilla
internodes (2-3
mm) between
florets
Lower
Glume

Upper
Glume

3.

GRASS ANATOMY Some Odds and Ends

[V. Max Brown]

Leaf sheaths are closed from base to middle or to near the top of the leaf sheath in the following;
1. Tribe Poeae (Genus Dactylis and some in the Genus Festuca)
2. Tribe Bromeae (Genus Bromus)
3. Tribe Melica (Genera Glyceria, Melica, Schizachne, and Pleuropogon).