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By Ian Ramjohn | May 4th 2009 11:00 AM | 6 comments | Print | E-mail | Track Comments
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More Tropical ecology notes articles

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About Ian Ramjohn

A Trinidadian in Oklahoma, I am a biology post-doc interested in tropical dry forests and island ecology. I also have a blog called Further Thoughts


... Full Bio

It's hard not to see the world through the lens of our own preconceptions and biases.  We tend to be more interested in other large mammals.  We're drawn to human-like qualities in pets.  But even the most benign insect is disturbingly alien when seen up close.  We also tend to use the familiar as a metaphor for understanding the unfamiliar.  Sometimes this gives us additional insight.  Other times, it leads us down the wrong path.

Most of us are familiar with mosses, even though we tend to use it as a catch-call term for any of the green 'gunk' that grows in damp areas (which tends to include not only mosses, but also cyanobacteria and lichens).  More botanically minded people will often combine mosses with liverworts and hornworts into a group known as the bryophytes.*
Closeup of mosses growing on a rock in Beacon Hill Park, Victoria, BC, Canada. Photo by Jesse Hickman, licensed under the Creative Commons Attribution-ShareAlike 2.5 License.

Colonisation of dry land was a major evolutionary challenge for plants (just as it was for animals).  Water provides buoyancy and support for the tissues of a plant.  Terrestrial plants must support their own tissues.  Land plants must deal with water loss and the ever-present risk of desiccation.  And finally, land plants have to deal with the fact that water and mineral nutrients are not even distributed in space; water and minerals are more abundant in the soil. 

Vascular plants - a group that includes flowering plants, conifers and ferns - have evolved a series of strategies to deal with these challenges.  Leaves are waterproofed by waxy cuticles.  Water is transported from the soil up to the top of the plant - even the tallest tree - through the xylem, which also doubles as a support system.

As traditionally defined, bryophytes were non-vascular plants  - plants that lack xylem and phloem. Without vascular tissue, they are unable to transport water and minerals very far from the soil surface.  This, coupled with their lack of tissues to prevent water loss, leaves them with a relatively limited set of options - they can only grow in moist environments, and they can never get very far from the group.

Most people would see this as a primitive condition - bryophytes have overcome some of the challenges their aquatic ancestors faced when they colonised the land, but only in a limited fashion.  This view, however, is a mistake.  Modern species - even "living fossils", which appear to have experienced little change in millions of years - are fully modern.  They are not primitive species that have somehow been sheltered from the realities of evolution.  They exist in and interact with communities of modern species.  Nonetheless, it's normal to think of "basal" taxa as having remained more similar to ancestral species, while more derived taxa have become more dissimilar.  But these assumptions can be misleading.

Bryophytes are unable able to conserve water; in seasonal environments it's normal to see shrivelled, desiccated mosses during dry periods.  Knowing that land plants originated from aquatic ancestors, it may seem reasonable to assume that this is a primitive condition, a failure, on the part of bryophytes to fully adapt to the challenges of life on land.  It would be a mistake to do so.  We shouldn't look at the inability to bryophytes to tolerate desiccation as an inferior solution to the problem of life on land.  Instead, it should be seen as an alternative solution to the problem.  Vascular plants have worked around the problem of desiccation by maximising water supply and minimising water loss.  Bryophytes, on the other hand, have evolved the ability to tolerate desiccation. 

There's a strong temptation to think of this as a "primitive" or "ancestral" condition, but in reality it provides options unavailable to vascular plants, like the ability to grow on rock surfaces or tree bark, places where there just isn't enough water available to supply the needs of vascular plants between wet periods.

*The traditional concept of the Bryophyta does not hold up under modern phylogenetic classification systems.  Liverworts are believed to be a sister group to all other land plants.  Mosses diverged next, with the final split between the hornworts and vascular plants coming later still.

Comments

Steve Davis's picture
You're right Ian, we do tend to label some plants as primitive, but I guess anything that works is a winner!

How the different Bryophytes are related to the vascular plants is still quite unclear, different studies come to different results.<?xml:namespace prefix = o ns = "urn:schemas-microsoft-com:office:office" />


For example (Nishiyama et al. 2004)&(Goremykin et al. 2005) gets the bryophytes as a monophyletic group and (Qiu et al. 2006) & (Groth-Malonek et al. 2005) gets the hornworts as the sister group to the vascular plants.


 


Mosses usually have some conducting tissue that is not so well developed as in vascular plant and even a weak cuticle. Polytricum is a genera with unusually well developed conducting tissue and the leaves moves to the stem when the plant gets dry. (Shaw 2000, Bryophyte biology)



The idea that some species can be more primitive is quite strange when al species has equal time to evolve and are "advanced" in different way f.ex. plants can syntesise loots of chemicals and animals has a complicated celd differential. 


Referenser


Qiu, Yin-Long et al. 2006.The deepest divergences in land plants inferred from phylogenomic evidence. PNAS vol. 103. pages 15511-15516


 


Goremykin, V.V.  and Hellwig, F.H.  2005. Evidence for the most basal split in land plants dividing bryophyte and tracheophyte lineages. Plant Systematics and Evolution 254 pages 93-103


 


Milena Groth-Malonek, Dagmar Pruchner, Felix Grewe & Volker Knoop. 2005. Ancestors of Trans-Splicing Mitochondrial Introns Support Serial Sister Group Relationships of Hornworts and Mosses with Vascular Plants. Molecular Biology and Evolution vol. 22(1).  pages 117-125


 


Tomoaki Nishiyama, Paul G. Wolf, Masanori Kugita, Robert B. Sinclair,


Mamoru Sugita, Chika Sugiura,  Tatsuya Wakasugi, Kyoji Yamada,


Koichi Yoshinaga, Kazuo Yamaguchi, Kunihiko Ueda & Mitsuyasu Hasebe 2004. Chloroplast Phylogeny Indicates that Bryophytes Are Monophyletic. Molecular Biology and Evolution vol. 21(10). 1813–1819



rholley's picture
Please don't start a politically correct pogrom against the word "primitive".  If one were to take a trip in imagination back to the late Devonian or early Carboniferous, might we not find mosses already clinging to rocks and enduring periods of dessication?  Away from the swampy coal-forests, conditions must have been tough for land plants.

I expect, though, that we are very short of fossils of the "bryophyta".  Even so, it is nice to think of them as "family firms" that were in existence when coal was still in the air.  Such a contrast to the vascular plants, which are dominated by "Johnny-come-latelys", while the clubmosses and horsetails are so greatly reduced from the magnificent forms that stood proud in the Carboniferous forests.

iramjohn's picture
"Politically correct pogrom"?  I see.  Are you really unable to express your opinions without spewing disgusting insults?  Is it really that difficult to be civil? 

Hank's picture
Please don't start a politically correct pogrom against the word "primitive".

... is what he wrote and, having read him for a while, I am betting he did not mean it in anything except the most light-hearted of ways.

rholley's picture
Ian,

No offence was meant: maybe I'm a little too much given to hyperbolic humour.

Anyway, I very much appreciate your article.  The "bryophytes" are greatly under-esteemed.

Here is Marchantia polymorpha, notable for its ability to spread through gemmae from the cups visible on the thallus (especially in garden centres) and to be the only visible live plant where a pot where the "main" plant has been killed by gross over-watering (image adapated from Wikimedia).



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