Fern Allies 

As for the ferns, the sporophyte is dominant while the gametophyte is short-lived. If it is generally true that ferns have big leaves, fern allies have small leaves or none at all. 

We are going to be quite selective in which groups we study and in what depth.


Club mosses or lycopods

While recent revisions have split the genus Lycopodium into several genera we will ignore this in this simple treatment. 


Lycopodium cernuum - a Caribbean species

Lycopodium spp. are essentially creeping plants with tiny leaves, termed microphylls, spirally arranged around the stem. Note how the leaves at the shoot tips look different. These are spore-bearing leaves called sporophylls
These sporophylls are tightly grouped into 
cones or strobili. 

(Some ferns do have spore-bearing and sterile leaves but grouping the sporophylls into a cone is quite different and distinct.) 


Biodisc photomicrograph
At left is an L.S. of a Lycopodium  cone. 

Note the sporangium in the axil of each sporophyll. 

Each sporangium is full of numerous tiny spores, similar in size to those of ferns. 

Incidentally, these spores were once used in fireworks. Their high oil content makes them burn readily. More recently they have been used to "dust" condoms.


The spores of Lycopodium germinate to produce a tiny, short-lived, peg-shaped gametophyte. This usually forms an association with a fungus and some species of Lycopodium have lost their photosynthetic capacity and rely on the fungus to provide carbohydrate! The gametophyte bears antheridia and archegonia and motile sperm cells swim to the eggs to fertilize these. The young sporophyte, as for ferns, develops initially as a parasite on the gametophyte. 

A = antheridia 
B = archegonia 
C = antherozoids

Check out the Lycopodium images at the University of Wisconsin

Extinct lycopods  



Lepidodendron  was a giant tree-like club moss. It towered to 45 m in height and flourished in the swamp forests of the Carboniferous. 


Another striking example of a tree-like, extinct lycopod is Sigillaria

Click here to learn more about fossil lycopods.


Spike mosses

 There is only one surviving genus, Selaginella

Selaginella serpens - a Caribbean native
There are about 700 Selaginella species world wide and several in the Caribbean. The sporophyte has a creeping habit and is dorsiventrally flattened. As in Lycopodium, there are separate sterile and fertile leaves and the spore-bearing leaves are grouped to form cones or strobili at the shoot tips.


In Selaginella the roots are typically borne on a rhizophore which extends from the stem.


Biodisc photomicrograph In the LS of the cone of Selaginella (left), you should be able to see that there are two kinds of spores: - 

microspores (small, many) 
megaspores (large, few) 

This is termed heterospory  and evolved several times in several groups of land plants. 

(Homospory is where there is only one kind of spore. Except for a few water ferns which are heterosporous the ferns are homosporous.) 

Click here to see this section in relation to a live plant.


The consequence of heterospory is separate male and female gametophytes;-  

 Each megasporophyll bears a megasporangium which by meoisis produces 4 megaspores which in turn form female gametophytes and female gametes. 
Each microsporophyll bears a microsporangium which by meoisis produces many microspores which in turn form male gametophytes and male gametes. 


In Selaginella, gametophytes develop within the actual spores, a phenomenon termed

Microspore split exposing sperm
 Each haploid microspore divides internally by mitosis to form ultimately an antheridium with 128 or 256 biflagellate sperm cells. These are released from the microspore wall and swim in surface water to megaspores . 
Cross section of megaspore showing archegonia at exposed top surface
Each haploid megaspore divides internally by free nuclear division to form a female gametophyte with archegonia opening to the spore surface. Sperm cells swim down the necks of these to fertilize eggs and produce the new diploid sporophyte generation. Click here to see this.


Three features distinguish Selaginella from Lycopodium;-  

Heterospory:  Selaginella  produces microspores and megaspores; Lycopodium is homosporous, producing only microspores. 
Endospory:  Selaginella microspores and megaspores develop their gametophytes within the actual walls of the spores; in Lycopodium the gametophyte grows out of the spore. 
Ligule: Each leaf in Selaginella (fertile and sterile) bears in its axil a tiny tongue-like structure called a ligule; these are not present in Lycopodium.  

You can check out more Selaginella images at the University of Wisconsin


Other fern allies

There are 3 other fern allies you should be aware of, although no detailed  knowledge of
these is required in this course.


The horsetails are recognizable by their erect, jointed stems ringed by microphylls and topped by cones. Only a single genus Equisetum survives today with a few temperate species but these plants were well represented in the swamp forests of the Carboniferous.  

Living Equisetum arvense, a herb Calamites, an extinct tree of the Carboniferous period
pterido-equisetum.jpg (17987 bytes)

pterido-calamites.jpg (37870 bytes) both of these beautiful  drawings are by Françoise Gantet

Equisetum_arv_sperm.JPEG (33951 bytes)
As for many other pteridophytes, the sperm cells of Equisetum are multiflagellate and very stream-lined.   



scanning electron micrographs by Karen Renzaglia


The Casuarina tree (Casuarina equisetifolia) is a flowering plant commonly cultivated in the Caribbean.  It gets it specific name from the resemblance of its branches to Equisetum. (equisetifolia = leaves like Equisetum

You can check out more Equisetum images at the University of Wisconsin 

Equisetum sp.


Whisk ferns   


Psilotum nudum  bearing sporangia.

We met Psilotum  when we looked at Rhynia, to which it bears a superficial resemblance. 

Rather than being living relatives of Rhynia, the whisk ferns are viewed by many experts as being  highly evolved relatives of the true ferns.

More on whisk ferns.



Only two genera are known, Isoetes and Stylites. They resemble clumps of grass-like leaves. 

Stylites  was only discovered in 1957 in the Andes. It is an a swamp-dwelling plant with no stomata, taking in CO2  through its roots! 

Isoetes is more widespread.


  Here is a photo of Isoetes in the flesh

The terms ferns and fern allies have served botanists well but be aware that phylogenies based on molecular studies are overturning these long held groupings. Recent studies (Pryer et al., 2004) suggest instead the following two phylogenetic groupings;-

   Monilophytes - all ferns, whisk ferns, horsetails

   Lycophytes - clubmosses, spikemosses, quillworts



Would you like to look at the lab for this part of the course? 
If so, click the button. 
Would you like to look at some sample questions on this part of the course? 
If so, click the button. 
We have now completed our look at the Fern Allies. 
Click the button to move on to the Evolution of Seed Plants. 

Click here for a good review site for ferns and fern allies 


C.M. Sean Carrington, 13  April 1998
last modified 15 November, 2012