Hydatellaceae: The Other Basal Angiosperms

Trithuria_submersa_-_Flickr_-_Kevin_Thiele.jpg

Though rather obscure to most of the world, the genus Trithuria has enjoyed somewhat of a celebrity status in recent years. A paper published in 2007 lifted this tiny group of minuscule aquatic plants out of their spot in Poales and granted them a place among the basal angiosperm lineage Nymphaeales. This was a huge move for such little plants. 

The genus Trithuria contains 12 species, the majority of which reside in Australia, however, two species, T. inconspicua and T. konkanensis, are native to New Zealand and India. They are all aquatic herbs and their diminutive size and inconspicuous appearance make them easy to miss. For quite some time these odd plants were considered to be a group of highly reduced monocots. Their original placement was in the family Centrolepidaceae. All of that changed in 2007.

Trithuria submersa DJD1631 Swedes Flat plants 2.jpg

Close inspection of Trithuria DNA told a much different story. These were not highly reduced monocots after all. Instead, multiple analyses revealed that Trithuria were actually members of the basal angiosperm lineage Nymphaeales. Together with the water lilies (Nymphaeaceae) and the fanworts (Cabombaceae), these plants are living representatives of some of the early days in flowering plant evolution. 

Of course, DNA analysis cannot stand on its own. The results of the new phylogeny had to be corroborated with anatomical evidence. Indeed, closer inspection of the anatomy of Trithuria revealed that these plants are truly distinct from members of Poales based on a series of features including furrowed pollen grains, inverted ovules, and abundant starchy seed storage tissues. Taken together, all of these lines of evidence warranted the construction of a new family - Hydatellaceae.

Trithuria_submersa_in_fruit.jpg

The 12 species of Trithuria are rather similar in their habits. Many live a largely submerged aquatic lifestyle in shallow estuarine habitats. As you may have guessed, individual plants look like tiny grass-like rosettes. Their small flower size has lent to some of their taxonomic confusion over the years. What was once thought of as individual flowers were revealed to be clusters or heads of highly reduced individual flowers. 

Reproduction for these plants seems like a tricky affair. Some have speculated that water plays a role but close inspections of at least one species revealed that very little pollen transfer takes place in this way. Wind is probably the most common way in which pollen from one plant finds its way to another, however, the reduced size of these flowers and their annual nature means there isn't much time and pollen to go around. It is likely that most of the 12 species of Trithuria are self-pollinated. This is probably quite useful considering the unpredictable nature of their aquatic habitats. It doesn't take much for these tiny aquatic herbs to establish new populations. In total, Trithuria stands as living proof that big things often come in small packages. 

Photo Credits: [1] [2] [3]

Further Reading: [1] [2] [3]

 

The Power of Leaves

When we think of the dominance of flowering plants on the landscape, we usually invoke the evolution of flowers and seed characteristics like endosperm and fruit. However, evolutionary adaptations in the structure of the angiosperm leaf may have been one of the most critical factors in the massive diversification that elevated them to their dominant position on the landscape today. 

Leaves are the primary organs used in water and gas exchange. They are the centers of photosynthesis, allowing plants to take energy from our closest star and turn it into food. To optimize this system, plants must balance water loss with transpiration in order to maximize their energy gain. This requires a complex plumbing system that can deliver water where it needs to be. It makes sense that plant physiology should maximize vein production, however, there are tradeoffs in doing so. Veins are not only costly to construct, they also displace valuable photosynthetic machinery. 

It appears that this is something that flowering plants do quite well. Because leaves fossilize with magnificent detail, researchers are able to look back in time through 400 million years of leaf evolution. What they found is quite incredible. There appears to be a consistent pattern in the vein densities between flowering and non-flowering plants. The densities found in angiosperm leaves both past and present are orders of magnitude higher than all non-flowering plants. These high densities are unique to flowering plants alone. 

This innovation in leaf physiology allowed flowering plants to maintain transpiration and carbon assimilation rates that are three and four times higher than those of non-flowering plants. This gives them a competitive edge across a multitude of different environments. The evolution of such dense vein structure also had major ramifications on the environment. 

The massive change in transpiration rates among the angiosperm lineage is likely to have completely changed the way water moved through the environment. These effects would be most extreme in tropical regions. Today, transpiration from tropical forests account for 30-50% of precipitation. A lot of this has to do with patterns in the intertropical convergence zone, which ensures that such humid conditions can be maintained. However, in areas outside of this zone such as in the Amazon, a high abundance of flowering plants with their increased rates of transpiration enhances the amount of rainfall and thus forms a sort of positive feedback.

Because precipitation is the single greatest factor in maintaining plant diversity in these regions, increases in rainfall due to angiosperm transpiration effectively helps to maintain such diversity. As angiosperms rose to dominance, this effect would have propagated throughout the ecosystems of the world. Plants really are the ultimate ecosystem engineers. 

Photo Credit: Bourassamr (Wikimedia Commons)

Further Reading: [1]

Anise: An Angiosperm Success Story

Illicium floridanum

Illicium floridanum

I must admit there are few flavors I loath more than anise (and consequently licorice and fennel). Regardless of the flavor, I nonetheless find myself enamored by their whorled seed capsules of star anise. In an attempt to reconcile my feelings towards anise in a culinary sense, I decided to get to know the plants that are responsible for it and I am so glad that I did. As it turns out, this group of small trees and shrubs offer us a glimpse at some of the earliest branchings on the angiosperm family tree.

We get star anise from the genus Illicium. Native to humid tropical understories, there are roughly 40 species scattered around southeast Asia, southeastern North America, the Caribbean, and parts of Mexico. Molecular as well as fossil evidence suggests this group diverged during the mid to late Cretaceous, not long after flowering plants came onto the scene. Indeed, along with Amborella and Nymphaeales, Illicium represent the three lineages that are sister to all other flowering plants alive today.

Illicium henryi

Illicium henryi

To call them primitive, however, would be a serious misnomer. Because they diverged so early on, these lineages represent serious success stories in flowering plant evolution. Instead, think of them as fruitful early experiments in angiosperm evolution. Illicium has characteristics that set it out as being sister to all other flowering plants. For instance, the vascular tissues more closely resemble those of gymnosperms than they do angiosperms. Also, like the other sister angiosperms, Illicium blur the line between the long standing categories of monocot and eudicot. As such, they are sometimes referred to as "paleoherbs." Another key diagnostic feature lies in their floral morphology.

They don't have what could be considered true petals or sepals. Instead, they have whorls of tepals, which start off sepal-like and gradually become more petal-like as you near the center of the flower. The stamens, which are laminar or leaf-like, are also arranged in a dense whorl surrounding a yet another whorl of fused carpels. Once fertilized, each carpel gives rise to a hard, glossy seed. As the carpels mature and begin to dry, the individual capsules get tighter and tighter until at some point the seed is pinched so hard that it is ejected from a slit in the fruit in projectile fashion.

Illicium verum

Illicium verum

Although this research will never rectify the taste of this spice, it nonetheless has given me a new found respect and sense of awe for this genus. To look upon the fruit of Illicium is to look at a biological structure that has stood the test of time. These plants are evolutionary successes that should be admired for their unique place in the story of flowering plant evolution.

Photo Credits: Scott Zona and Tim Waters

Further Reading: [1]

How Leaf Veins Changed the World

When we think of the dominance of flowering plants on the landscape, we usually invoke the evolution of flowers and seed characteristics such as an endosperm and fruit. However, evolutionary adaptations in the structure of the angiosperm leaf may have been one of the critical factors in the massive diversification that elevated them to their dominant position on the landscape today.

Leaves are the primary organs used in water and gas exchange. They are the centers of photosynthesis, allowing plants to take energy from our closest star and turn it into food. To optimize this system, plants must balance water loss with transpiration in order to maximize their energy gain. This requires a complex plumbing system that can deliver water where it needs to be. It makes sense that plant physiology should maximize vein production, however, there are tradeoffs in doing so. Veins are not only costly to construct, they also displace valuable photosynthetic machinery.

It appears that this is something that flowering plants do quite well. Because leaves fossilize with magnificent detail, researchers are able to look back in time through 400 million years of leaf evolution. What they found is quite incredible. There appears to be a consistent pattern in the vein densities between flowering and non-flowering plants. The densities found in angiosperm leaves both past and present are orders of magnitude higher than all non-flowering plants. These high densities are unique to flowering plants alone.

This innovation in leaf physiology allowed flowering plants to maintain transpiration and carbon assimilation rates that are three and four times higher than those of non-flowering plants. This gives them a competitive edge across a multitude of different environments. The evolution of such dense vein structure also had major ramifications on the environment.

This massive change in transpiration rates among the angiosperm lineage is likely to have completely changed the way water moved through the environment. These effects would be most extreme in tropical regions. Today, transpiration from tropical forests account for 30-50% of precipitation. A lot of this has to do with patterns in the intertropical convergence zone, which ensures that such humid conditions can be maintained. However, in areas outside of this zone such as in the Amazon, a high abundance of flowering plants with their increased rates of transpiration enhances the amount of rainfall and thus forms a sort of positive feedback. Because precipitation is the single greatest factor in maintaining plant diversity in these regions, increases in rainfall due to angiosperm transpiration effectively helps to maintain such diversity. As angiosperms rose to dominance, this effect would have propagated throughout the ecosystems of the world.

Photo Credit: Bourassamr (Wikimedia Commons)

Further Reading:
http://rspb.royalsocietypublishing.org/content/276/1663/1771

Cretaceous Seeds Shine Light on the Evolution of Flowering Plants

What you are looking at here are some of the earliest fossil remains of flowering plants. These seeds were preserved in Cretaceous sediments dating back some 125–110 million years ago. Fossil evidence dating to the early days of the angiosperm lineage is scant, which makes these fossils all the more spectacular. Thanks to a large collaborative effort, Dr. Else Marie Friis is shining light on the evolution of seeds.

Finding these fossils is not a matter of seeing them with the naked eye. These seeds are tiny, ranging from half a millimeter up to 2 millimeters in length. They were discovered using an advanced form of X-ray microscopy. The advantage of this technique is not only that it is nondestructive but it also allows researchers to investigate the internal structures of the seeds that would otherwise be impossible to see. Their preservation is mind blowingly delicate, allowing researchers to see minute details of the embryo and even subcellular structures like nuclei. 

Dr. Friis' team was able to look at over 250 fossil seeds from 75 different taxa. They were able to make 3D models of the embryos, allowing for more detailed studies than ever before. For some of the fossils, the detail was such that they were able to match them to extant lineages of flowering plants. For others, this technique is allowing for better reclassification of now extinct species. 

By far the most exciting part about these fossils are what they can tell us about the ecology of early flowering plants. In all instances, the embryos within the seeds were small, immature, and dormant. This suggests that seed dormancy is a fundamental trait of flowering plants. What's more, this lends support to the hypothesis that angiosperms first evolved as opportunistic, early successional colonizers. Seed dormancy allows flowering plants to wait out the bad times until favorable environmental conditions allowed for germination and seedling establishment. 

Photo Credit: Dr. Else Marie Friis

Further Reading:
http://www.nature.com/nature/journal/v528/n7583/full/nature16441.html

Aquatic Angiosperm: A Cretaceous Origin?

It would seem that yet another piece of the evolutionary puzzle that are flowering plants has been found. I have discussed the paleontological debate centered around the angiosperm lineage in the past (http://bit.ly/1S6WLkf), and I don't think the recent news will put any of it to rest. However, I do think it serves to expand our limited view into the history of flowering plant evolution.

Meet Montsechia vidalii, an extinct species that offers tantalizing evidence that flowering plants were kicking around some 130–125 million years ago, during the early days of the Cretaceous. It is by no means showy and I myself would have a hard time distinguishing its reproductive structures as flowers yet that is indeed what they are thought to be. Detailed (and I mean detailed) analyses of over 1,000 fossilized specimens reveals that the seeds are enclosed in tissue, a true hallmark of the angiosperm lineage.

On top of this feature, the fossils also offer clues to the kind of habitat Montsechia would have been found in. As it turns out, this was an aquatic species. The flowers, instead of poking above the water, would have remained submerged. An opening at the top of each flower would have allowed pollen to float inside for fertilization. Another interesting feature of Montsechia is that it had no roots. Instead, it likely floated around in shallow water.

This is all very similar to another group of extant aquatic flowering plants in the genus Ceratophyllum (often called hornworts or coon's tail). Based on such morphological evidence, it has been agreed that these two groups represent early stem lineages of the angiosperm tree. Coupled with what we now know about the habitat of Archaefructus (http://bit.ly/1S6WLkf), it is becoming evident that the evolution of flowers may have happened in and around water. This in turn brings up many more questions regarding the selective pressures that led to flowers.

What is even more amazing is that these fossils are by no means recent discoveries. They were part of a collection that was excavated in Spain over 100 years ago. Discoveries like this happen all the time. Someone finds a interesting set of fossils that are then stored away on a dark shelf in the bowels of a museum only to be rediscovered decades or even centuries later.

All in all I think this discovery lends credence to the idea that flowering plants are a bit older than we like to think. Also, one should be wary of anyone claiming to have found "the first flower." The idea that there could be a fossil out there that depicts the first anything is flawed a leads to a lot of confusion. Instead, fossils like these represent snapshots in the continuum that is evolution. Each new discovery reveals a little bit more about the evolution of that lineage. We will never find the first flower but we will continue to refine our understanding of life on this planet.

Photo Credits: Bernard Gomeza, Véronique Daviero-Gomeza, Clément Coiffardb, Carles Martín-Closasc, David L. Dilcherd, and O. Sanisidro,

Further Reading:
http://www.pnas.org/content/112/35/10985.abstract