Club Mosses, Ferns & Horsetails:

the Seed-free Vascular Plants

Vascular Plants - a quick review

Two unrelated groups within cryptogams seed free vascular
plants are recognized as phyla:

1. Lycopodiophyta : lycopods

2. Polypodiophyta: ferns, horsetails, and whisk ferns

Vascular Plants - a quick review

Why were the seed-free plants grouped together?

They produce free spores, the principal dispersal units, via meiosis.
Spore: a reproductive cell, capable of developing into an adult
without fusion with another cell.

spores

Vascular Plants - a quick review

Why were the seed-free plants grouped together?

Spores develop within a sporangium (pl. sporangia)

sporangium

spores

Vascular Plants - a quick review

Why were the seed-free plants grouped together?

Spores germinate and develop into gametophytes that exist
independently of the spore-producing plants. The gametophytes
(haploid, n) tend to be inconspicuous and short-lived.

sporangium

gametophyte

spores

Vascular Plants - a quick review

Why were the seed-free plants grouped together?

Like all plants, seed-free plants produce two kinds of gametes in their
gametophytes: sperm and egg that unite to form a zygote (2n or
diploid) via fertilization!
sporangium

zygote

gametophyte

spores

Vascular Plants - a quick review

Why were the seed-free plants grouped together?

The sporophyte (2n) develops from the zygote and is more
conspicuous, usually perennial and lives for an indefinite period!
sporangium

sporophyte

zygote

gametophyte

spores

Wisconsin Seed-free Plants

The best website to identify and see images of Wisconsin s seedfree plants is Gary Fewless at UW-Green Bay; links provided
below

Key to Ferns and Fern Allies of Wisconsin

List of Pteridophytes of Wisconsin
Glossary of Fern Terminology
also:
Michigan Online Flora

Wisconsin Seed-free Plants

The best manual to identify lycopods
and ferns is Ferns of Northeastern
and Central North America (2nd ed.)
in the Peterson Field Guides

Warning:



Families and genera (and thus
species names) are changing quickly
in the seed-free plants

Phylum Lycopodiophyta
club mosses, spike mosses, quillworts!

Leaves microphylls: generally small, simple,

one-veined leaves. Microphylls evolved by the
process of enation and vascularization. An
enation is a veinless, lateral protrubence on
the stem. When the enation becomes
vascularized it is termed a microphyll.!

Sporangia: the spore producers on the

sporophytes are located singly on the upper
surfaces or in axils of the bracts of a cone or of
green leaves!

Phylum Lycopodiophyta!

Carboniferous forest from Illinois

This group arose in the Later Silurian about 420 million years ago and
was dominant in the Carboniferous and used to be much more
diverse.!

Phylum Lycopodiophyta!
Lycopodiaceae - club mosses

15 genera and about 375 species



Lycopodium now split into several genera



cosmopolitan distribution, most diverse in tropics

Evergreen, stems elongate and dichotomously
branching. Leaves often densely covering the stem.

Oily compounds in the cell walls ignite rapidly into
a flash of light and were used by magicians and
sorcerers in the Middle Ages. More recently they
were used as a flash early in photography and in
experimental photocopying machines. Many
species over-collected for Christmas wreaths.

Diphasiastrum complanatum

Ground cedar, crowfoot

Phylum Lycopodiophyta!
Lycopodiaceae - club mosses

Homosporous: same spore - one kind of spore

produced; cones terete (rounded)

Phylum Lycopodiophyta!
Lycopodiaceae - club mosses

other examples

Dendrolycopodium obscurum

Ground pine

Phylum Lycopodiophyta!
Lycopodiaceae - club mosses

other examples

Lycopodiella inundata

Bog club moss

Phylum Lycopodiophyta!
Lycopodiaceae - club mosses

other examples

Huperzia lucidula

Shiny club moss

Alkaloid for Alzheimers?

Phylum Lycopodiophyta!
Huperzia lucidula

Shiny club moss

Polytrichum

forest mosses

Phylum Lycopodiophyta!
Selaginellaceae - spike mosses

1 genus and about 750 species



Mainly tropical family with some species
extending into arctic regions of both hemispheres

Leaves spirally arranged and often 4-ranked on the
secondary and ultimate branches.



Spores borne in or near the axils of well-differentiated
sprophylls, usually on 4 sided stroboli.

Phylum Lycopodiophyta!
Selaginellaceae - spike mosses

1 genus and about 750 species



Mainly tropical family with some species
extending into arctic regions of both hemispheres

Leaves spirally arranged and often 4-ranked on the
secondary and ultimate branches.



Spores borne in or near the axils of well-differentiated
sprophylls, usually on 4 sided stroboli.




Heterosporous: different spores !
!

Unlike Lycopodiaceae, Selaginellaceae are

heterosporous with different types of spores:
microspores and megaspore (not related to size) the
micro give rise to male gametophytes and the mega
give rise to the female gametophytes.

Phylum Lycopodiophyta!
Selaginellaceae - spike mosses

Heterosporous:

different spores !

Unlike Lycopodiaceae, Selaginellaceae are

heterosporous with different types of spores:
microspores and megaspore (not related to size) the
micro give rise to male gametophytes and the mega
give rise to the female gametophytes.

Phylum Lycopodiophyta!
Selaginellaceae - spike mosses

Selaginella eclipes

meadow spike moss

Phylum Lycopodiophyta!
Selaginellaceae - spike mosses

Selaginella ruprestis

Rock spike moss

Selaginella selaginoides

Northern spike moss

Endangered!

Phylum Lycopodiophyta!
Isoetaceae - quillworts

1 genus and about 150 species; worldwide; 2 spp.
in WI - Isoetes echinospora most common



Aquatic or semi-aquatic plants with corm-like
stems with secondary growth. Absorb carbon
through their roots.

Leaves with ligules and quite long (15+ cm) for
microphylls.

I. butleri not native

Isoetes echinospora

Phylum Lycopodiophyta!
Isoetaceae - quillworts

Heterosporous plants with sporangia borne at the base of sporophylls which are
similar to vegetative leaves. Megasporangia + microsporangia

Phylum Polypodiophyta - ferns & horsetails!

Equisetaceae - horsetails, scouring rushes
[previously placed in own phylum - Equisetophyta]

1 genus, Equisetum, 15 species with a

cosmopolitan distribution except for Australia or
New Zealand [9 species in Wisconsin]

Plants are primarily colonizers of unforested areas,
lake margins and wetlands

Shoots monomorphic or dimorphic (see right).



Often highly branched (horsetails) appearing
like leaves. Others not branched (scouring
rushes).

Internodes with conspicuous vertical ridges; jointed

stems; stems hollow (both important taxonomic
features for keying species)

Phylum Polypodiophyta!
Equisetaceae - horsetails, scouring rushes

Leaves in whorls; leaves one veined,
verticillate, united to form a sheath
around the stem; these leaves are probably
reduced megaphylls: Megaphylls are
larger than microphylls have a blade that
has a complex system of veins.

megaphyll

sheath

branch

Megaphyll theory is supported by

placement of Equisetum among
primitive ferns using DNA.

Phylum Polypodiophyta!
Equisetaceae - horsetails, scouring rushes

Sporangia clustered terminally in cones
composed of polygonal, umbrella-like
structures with sporangia beneath.

Phylum Polypodiophyta!
Equisetaceae - horsetails, scouring rushes

Horsetails are homosporous, have green spores with
hygroscopic elators, and form conspicuous green
gametophytes

Phylum Polypodiophyta!
Equisetaceae - some examples of native horsetails & scouring rushes

Equisetum arvense

Field horsetail

Equisetum sylvaticum

Woodland horsetail

Phylum Polypodiophyta!
Equisetaceae - some examples of native horsetails & scouring rushes

Equisetum laevigatum

Smooth scouring rush

Equisetum scirpoides

Dwarf horsetail

Phylum Polypodiophyta
the true ferns!
Most diverse of the vascular cryptogams, both in
species number (ca. 11,000 worldwide) and form
(small aquatics to trees).



Like the horsetails (which may be thought of as
specialized ferns), leaves are megaphylls; blade is
called a frond and the petioles as stipes.

Phylum Polypodiophyta
ferns!
Circinate vernation: vernation is the arrangement of folded leaves in a bud, forming a
crozier or fiddlehead, i.e. coiled or rolled up at the tip and unfolding lengthwise when
emerging - due to auxin and differential growth of tissue. Protects young bud.

Phylum Polypodiophyta
ferns!
Sporangia borne on the margin or the lower surface of the leaf; often grouped in sori
(pl.) sorus (sing.); a sorus may be protected by a flap-like structure called the
indusium - an important feature for identification of ferns due to its remarkable
variation in size, shape, and position.

Phylum Polypodiophyta
ferns!
The sprorangium is often associated with an annulus a cluster or row of cells with
thick walls that open the sporangium and catapult the spores into the air.

Ferns can be either homosporous
or heterosporous.

annulus

Phylum Polypodiophyta
ferns!

Phylum Polypodiophyta!
There is major disagreement on what are the fern families, but 30 are generally
recognized worldwide. In Wisconsin there are 9 families.



Characters used to circumscribe families often relate to cryptic features such as
the presence or absence of annulus and its orientation, the length and diameter
(number of cells) of the sporangial stalk, spore shape, stem and petiole crosssection anatomy. The primitive ferns include horsetails and whiskferns.

Phylum Polypodiophyta!
Ophioglossaceae - adder s tongue family

Primitive ferns; 2 genera in Wisconsin:
Ophioglossum and Botrychium (now 3 genera)

Sporangia are on an erect axis

with a green blade attached below

the sporangia bearing part

Ophioglossum pusillum

Adder s tongue fern

Phylum Polypodiophyta!
Ophioglossaceae - adder s tongue family

Primitive ferns; 4 genera in Wisconsin:
Ophioglossum and Botrychium (now 3 genera)

Botrychium mormo

Goblin fern

Botrychium lunaria

Moonwort fern

Botrypus virginianum

Rattlesnake fern

Phylum Polypodiophyta!
Osmundaceae - royal fern family

Primitive ferns;

1 genus in Wisconsin: Osmunda

Fertile and sterile
leaves dimorphic and
the fertile portion is
non-green at maturity.
Sporangia not
clustered in sori, but
naked.

Phylum Polypodiophyta!
Osmundaceae - royal fern family

3 species of Osmunda in Wisconsin;
easily separated by position of fertile
portions

Osmunda cinnamomea

Cinnamon fern

All wide-spread &

Bot 401 exam plant!

Osmunda claytoniana

Interrupted fern

Osmunda regalis

Royal fern

Phylum Polypodiophyta!
Dennstaedtiaceae - bracken family

2 genera in Wisconsin. Includes one of
the most widespread of all vascular
plants.

Clonal with rhizome; large compound

leaves. Ubiquitous in Wisconsin.

Marginal sori with no
indusia but with revolute
(rolled over) leaf edge
protecting sori.

Pteridium aquilinum

Bracken fern

Phylum Polypodiophyta!
Pteridaceae - maidenhair fern family

4 genera in Wisconsin

Sori that lack indusia or are protected

by a reflexed or revolute margins

Distinctively compound
frond with dark purple
stipe and rachis

Adiatum pedatum

Maidenhair fern

Phylum Polypodiophyta!
Aspleniaceae - spleenwort family

1 genus in Wisconsin. Defined by linear
or kidney shaped sori.

Asplenium viride

(A. trichomanes-ramosum)

Green spleenwort

Asplenium platyneuron

Ebony spleenwort

Phylum Polypodiophyta!
Onocleaceae- sensitive fern family

2 genera in Wisconsin (also Matteuccia ostrich
fern). Sensitive ferns has dimorphic fronds. Sterile
frond pinnately lobed. Fertile frond turning black.

Onoclea sensibilis

Sensitive fern

Phylum Polypodiophyta!
Dryopteridaceae - woodfern family

Large and diverse group of ferns; often broadly defined to include
other smaller families. 2 genera and 13 species in Wisconsin.

Polystichum acrostichoides

Christmas fern

Dryopteris intermedia

Shield fern

Phylum Polypodiophyta!
Polypodiaceae - rockcap family

40 genera and over 500 species in
tropics and subtropics, but a single
species in Wisconsin.

Distinctive fronds: simple, pinnately

lobed; leathery. Colonizes bare rock.

Sori in two rows on each
lobe.

Polypodium virginianum

Rockcap fern

Phylum Polypodiophyta!
Salviniaceae (Azollaceae) - mosquito fern family

Western & SE coastal elements

2 species in Wisconsin. Floating aquatic, reduced ferns with

2 ranked leaves, each 2 lobed. Symbiotic relationship with
N2 fixing blue green bacteria Anabaena azollae.
Heterosporous!

Azolla mexicana & A. caroliniana

Floating fern