The Rise and Fall of the Scale Trees

Photo by Ghedoghedo licensed under CC BY-SA 3.0

If I had a time machine, the first place I would visit would be the Carboniferous. Spanning from 358.9 to 298.9 million years ago, this was a strange time in Earth’s history. The continents were jumbled together into two great landmasses - Laurasia to the north and Gondwana to the south and the equatorial regions were dominated by humid, tropical swamps. To explore these swamps would be to explore one of the most alien landscapes this world has ever known.

The Carboniferous was the heyday for early land plants. Giant lycopods, ferns, and horsetails formed the backbone of terrestrial ecosystems. By far the most abundant plants during these times were a group of giant, tree-like lycopsids known as the scale trees. Scale trees collectively make up the extinct genus Lepidodendron and despite constantly being compared to modern day club mosses (Lycopodiopsida), experts believe they were more closely related to the quillworts (Isoetopsida).

Carboniferous coal swamp reconstruction dating back to the 1800’s

It is hard to say for sure just how many species of scale tree there were. Early on, each fragmentary fossil was given its own unique taxonomic classification; a branch was considered to be one species while a root fragment was considered to be another, and juvenile tree fossils were classified differently than adults. As more complete specimens were unearthed, a better picture of scale tree diversity started to emerge. Today I can find references to anywhere between 4 and 13 named species of scale tree and surely more await discovery. What we can say for sure is that scale tree biology was bizarre.

The name “scale tree” stems from the fossilized remains of their bark, which resembles reptile skin more than it does anything botanical. Fossilized trunk and stem casts are adorned with diamond shaped impressions arranged in rows of ascending spirals. These are not scales, of course, but rather they are leaf scars. In life, scale trees were adorned with long, needle-like leaves, each with a single vein for plumbing. Before they started branching, young trees would have resembled a bushy, green bottle brush.

As scale trees grew, it is likely that they shed their lower leaves, which left behind the characteristic diamond patterns that make their fossils so recognizable. How these plants achieved growth is rather fascinating. Scale tree cambium was unifacial, meaning it only produced cells towards its interior, not in both directions as we see in modern trees. As such, only secondary xylem was produced. Overall, scale trees would not have been very woody plants. Most of the interior of the trunk and stems was comprised of a spongy cortical meristem. Because of this, the structural integrity of the plant relied on the thick outer “bark.” Many paleobotanists believe that this anatomical quirk made scale trees vulnerable to high winds.

Scale trees were anchored into their peaty substrate by rather peculiar roots. Originally described as a separate species, the roots of these trees still retain their species name. Paleobotanists refer to them as “stigmaria” and they were unlike most roots we encounter today. Stigmaria were large, limb-like structures that branched dichotomously in the soil. Each main branch was covered in tiny spots that were also arranged in rows of ascending spirals. At each spot, a rootlet would have grown outward, likely partnering with mycorrhizal fungi in search of water and nutrients.

A preserved Lepidodendron stump

Eventually scale trees would reach a height in which branching began. Their tree-like canopy was also the result of dichotomous branching of each new stem. Amazingly, the scale tree canopy reached staggering heights. Some specimens have been found that were an estimated 100 ft (30 m) tall! It was once thought that scale trees reached these lofty heights in as little as 10 to 15 years, which is absolutely bonkers to think about. However, more recent estimates have cast doubt on these numbers. The authors of one paper suggest that there is no biological mechanism available that could explain such rapid growth rates, concluding that the life span of a typical scale tree was more likely measured in centuries rather than years.

Regardless of how long it took them to reach such heights, they nonetheless would have been impressive sites. Remarkably, enough of these trees have been preserved in situ that we can actually get a sense for how these swampy habitats would have been structured. Whenever preserved stumps have been found, paleobotanists remark on the density of their stems. Scale trees did not seem to suffer much from overcrowding.

The fact that they spent most of their life as a single, unbranched stem may have allowed for more success in such dense situations. In fact, those that have been lucky enough to explore these fossilized forests often comment on how similar their structure seems compared to modern day cypress swamps. It appears that warm, water-logged conditions present similar selection pressures today as they did 350+ million years ago.

Like all living things, scale trees eventually had to reproduce. From the tips of their dichotomosly branching stems emerged spore-bearing cones. The fact that they emerge from the growing tips of the branches suggests that each scale tree only got one shot at reproduction. Again, analyses of some fossilized scale tree forests suggests that these plants were monocarpic, meaning each plant died after a single reproductive event. In fact, fossilized remains of a scale tree forest in Illinois suggests that mass reproductive events may have been the standard for at least some species. Scale trees would all have established at around the same time, grown up together, and then reproduced and died en masse. Their death would have cleared the way for their developing offspring. What an experience that must have been for any insect flying around these ancient swamps.

The fossilized strobilus of a Lepidodendron. Photo by Verisimilus T licensed under the GNU Free Documentation License.

Compared to modern day angiosperms, the habits of the various scale trees may seem a bit inefficient. Nonetheless, this was an extremely successful lineage of plants. Scale trees were the dominant players of the warm, humid, equatorial swamps. However, their dominance on the landscape may have actually been their downfall. In fact, scale trees may have helped bring about an ice age that marked the end of the Carboniferous.

You see, while plants were busy experimenting with building ever taller, more complex anatomies using compounds such as cellulose and lignin, the fungal communities of that time had not yet figured out how to digest them. As these trees grew into 100 ft monsters and died, more and more carbon was being tied up in plant tissues that simply weren’t decomposing. This lack of decomposition is why we humans have had so much Carboniferous coal available to us. It also meant that tons of CO2, a potent greenhouse gas, were being pulled out of the atmosphere millennia after millennia.

A fossilized root or “stigmaria”. Photo by Verisimilus T licensed under CC BY-SA 3.0

As atmospheric CO2 levels plummeted and continents continued to shift, the climate was growing more and more seasonal. This was bad news for the scale trees. All evidence suggests that they were not capable of keeping up with the changes that they themselves had a big part in bringing about. By the end of the Carboniferous, Earth had dipped into an ice age. Earth’s new climate regime appeared to be too much for the scale trees to handle and they were driven to extinction. The world they left behind was primed and ready for new players. The Permian would see a whole new set of plants take over the land and would set the stage for even more terrestrial life to explode onto the scene.

It is amazing to think that we owe much of our industrialized society to scale trees whose leaves captured CO2 and turned it into usable carbon so many millions of years ago. It seems oddly fitting that, thanks to us, scale trees are once again changing Earth’s climate. As we continue to pump Carboniferous CO2 into our atmosphere, one must stop to ask themselves which dominant organisms are most at risk from all of this recent climate change?

Photo Credits: [1] [2] [4] [5] [6] [7]

Further Reading: [1] [2] [3] [4] [5] [6] [7] [8] [9] [10] [11] [12] [13] [14]

Fossils Shine Light On the History of Gall-Making Wasps

M J Richardson / Common spangle galls / CC BY-SA 2.0

We can learn a lot about life on Earth from the fossil record. I am always amazed by the degree of scrutiny involved in collecting data from these preserved remains. Take, for instance, the case of gall-making wasp fossils found in western North America. A small collection of fossilized oak leaves is giving researchers insights into the evolutionary history of oaks and the gall-making wasps they host.

Oaks interact with a bewildering array of insects. Many of these are gall-making wasps in the family Cynipidae. Dozens of different wasp species can be found on a single oak tree. Female wasps lay their eggs inside developing oak tissues and the larvae release hormones and other chemicals that cause galls to form. Galls are essentially edible nursery chambers. Other than their bizarre shapes and colors, the compounds released by the wasp larvae reduce the chemical defenses of the oak and increase the relative nutrition of the tissues themselves. Often, these relationships are precise, with specific wasp species preferring specific oak species. But when did these relationships arise? Why are oaks so popular? What can fossil evidence tell us about this incredible relationship?

Photo by Beentree licensed under CC BY-SA 4.0

Though scant, the little fossil evidence of oak galls can tell us a lot. For starters, we know that gall-making wasps whose larvae produce structures similar to that of the Cynipids have been around since at least the late Cretaceous, some 100 million years ago. However, it is hard to say for sure exactly who made these galls and exactly what taxonomic affinity the host plant belongs to. More conclusive Cynipid gall fossils appear again in the Eocene and continue to pop up in the fossil record throughout the Oligocene and well into the Miocene (33 - 23 million years ago).

Miocene aged fossils are where things get a little bit more conclusive. Fossil beds located in the western United States have turned up fossilized oak leaves complete with Cynipid galls. The similarity of these galls to those of some present day species is incredible. It demonstrates that these relationships arose early on and have continued to diversify ever since. What's more, thanks to the degree of preservation in these fossil beds, researchers are able to make some greater conclusions about why gall-making wasps and oaks seem to be so intertwined.

Holotype of Antronoides cyanomontanus galls on fossilized leaves of Quercus simulata. 1) Impression of the abaxial surface of the leaf, showing the galls extending into the matrix. 2) Galls showing close association with secondary veins. 3) Gall showing the impression of rim-like base partially straddling the secondary vein. 4) Close-up of gall attached at margin extending down into the matrix. 5) Gall uncovered revealing spindle-shaped morphology.

1) Xanthoteras clavuloides galls on fossilized Quercus lobata, showing gall attached to secondary vein. Specimen in California Academy of Sciences Entomology collection, San Francisco. 2) Two galls of attached to a secondary vein showing overlap of their bases. Specimen in California Academy of Sciences Entomology Collection, San Francisco. 3) Three galls collected from leaf of California Quercus lobata showing clavate shape and expanded, ring-like base. 4) Gall showing the annulate or ribbed aspect of the base, which is similar to bases of Antronoides cyanomontanus and A. polygonalis. 5) Galls showing clavate shape, pilose and nonpilose surfaces, and bases.

Gall-making wasps seem to diversify at a much faster rate in xeric climates. The fossil records during this time show that mesic tree speciess were gradually being replaced by more xeric species like oaks. Wasps seem to prefer these drier environments and the thought is that such preferences have to do with disease and parasite loads.

Again, galls a large collections of nutrient-rich tissues that are low in defense compounds. Coupled with the juicy grub at their center, it stands to reason that galls make excellent sites of infection for fungi and other parasites. By living in drier habitats, it is believed that gall-making wasps are able to escape these environmental pressures that would otherwise plague them in wetter habitats. The fossil evidence appears to support this hypothesis and today we see similar patterns. White oaks are especially drought tolerant and its this group of oaks that host the highest diversity of gall-making wasps.

Photo Credits: [1] [2] [3] [4]

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

The First Trees Ripped Themselves Apart To Grow

Illustration by Falconaumanni licensed under CC BY-SA 3.0

A new set of fossil discoveries show that the evolutionary arms race that are forests started with plants that literally had to rip themselves apart in their battle for the canopy. The first forests on this planet arose some 385 million years ago and were unlike anything we know today. They consisted of a clade of trees known scientifically as Cladoxylopsids, which have no living representatives in these modern times. How these trees lived and grew has remained a mystery since their fossilized trunks were first discovered but a new set of fossils from China reveals that these trees were unique in more ways than one.

Laying eyes on a full grown Cladoxylopsid would be a strange experience to say the least. Their oddly swollen base would gradually taper up a trunk that stretched some 10 to 12 meters (~30 - 40 feet) into a canopy of its relatives. They had no leaves either. Instead, their photosynthetic organs consisted of branch-like growths that were covered in twig-like projections. Whereas most fossils revealed great detail about their outward appearance, we have largely been in the dark on what their internal anatomy was like. Excitingly, a set of exquisitely preserved fossils from Xinjiang, China has changed that. What they reveal about these early trees is quite remarkable.

[SOURCE]

As it turns out, the trunks of these early trees were hollow. Unlike the trees we know today, whose xylem expands in concentric rings and forms a solid trunk, the trunk of Cladoxylopsid was made up of strands of xylem connected by a network of softer tissues. Each of these strands was like a mini tree in and of itself. Each strand formed its own concentric rings that gradually increased the size of the trunk. However, this gradual expansion did not appear to be a gentle process.

As these strands increased in size, the trunk would grow larger and larger. In doing so, the tissues connecting the strands were pulled tighter and tighter. Eventually they would tear under the strain. They would gradually repair themselves over time but the effect on the trunk was quite remarkable. In effect, the base of the tree would literally collapse in on itself in a controlled manner. You could say that older Cladoxylopsids developed a bit of a muffin top at their base. 

A cross section of a Cladoxylopsid trunk showing the hollow center, individual xylem strands, and the network of connective tissues. [SOURCE]

Although this seems very detrimental, the overall structure of the tree would have been sturdy. The authors liken this to the design of the Eiffel tower. Indeed, a hollow cylinder is actually stronger than a solid one of the same dimensions. When looked at in the context of all other trees, this form of growth is truly unique. No other trees are constructed in such a manner.

The authors speculate that this form of growth may be why these trees eventually went extinct. It would have taken a lot of energy to grow in that manner. It is possible that, as more efficient forms of growth were evolving, the Cladoxylopsids may not have been able to compete. It is anyone's guess at this point but this certainly offers a window back into the early days of tree growth. It also shows that there has always been more than one way to grow a tree.

LEARN MORE ABOUT THESE TREES AND THE FORESTS THEY MADE IN EPISODE 253 OF THE IN DEFENSE OF PLANTS PODCAST.

Photo Credits: [1] [2]

Further Reading: [1]

Ferns Unchanged

Ferns are old. Arising during the late Devonian period, some 360 million years ago, ferns once dominated the land. These ancient ferns were a bit different than the ferns we know today. It wasn't until roughly 145 million years ago, during the late Cretaceous period, that many extant fern families started to appear. However, a recent fossil discovery shows that at least one familiar fern was hanging out with dinosaurs as far back as 180 million years ago!

A team of scientists in Sweden recently unearthed an exquisitely preserve fossil of a fern from some early Jurassic deposits. Usually the fossilization process does not preserve very fine details, especially not at the cellular level, but that is not the case for this fossil. Falling into volcanic hydrothermal brine, the fern quickly mineralized. The speed at which the tissues of the fern were replaced by minerals preserved details that paleontologists usually only dream about. Clearly visible in the fossilized stem are subcellular structures like nuclei and even chromosomes in various stages of cell division!

 

A) Section of the fossil rhizome. B-J) Exquisitely preserve cellular details [SOURCE]

Using sophisticated microscopy techniques, the team was able to analyze the properties of the nuclei undergoing division. What they discovered is simply amazing. The number of chromosomes as well as other properties of the DNA matched a fern that is quite common in eastern North America and Asia today. This fossilized fern, as far as the team can tell, is a close relative of the cinnamon fern (Osmundastrum cinnamomeum), placing it in the royal fern family (Osmundaceae). Based on the fossil evidence, relatives of these ferns were not only around during the early Jurassic, they have remained virtually unchanged for 180 million years. Talk about living fossils!

Further Reading: [1] [2]

Paleo Pinus

Photo Credit: Howard Falcon-Lang, Royal Holloway University of London

What you are looking at here is the oldest fossil evidence of the genus Pinus. Now, conifers have been around a long time. I mean really long. Recognizable members of this group first came onto the scene sometime during the late Triassic, some 235 million years ago. Today, one of the most species-rich genera of conifers are those in the genus Pinus. They dominate northern hemisphere forests and can be found growing in dry soils throughout the globe. For such a commonly encountered group, their origins have remained a bit of a mystery. 

The fossil was discovered in Nova Scotia, Canada. Unlike the rocky fossils we normally think of, this fossil was preserved as charcoal, undoubtedly thanks to a forest fire. The degree of preservation in this charcoal specimen is astounding and provides ample opportunity for close investigation. 

I mentioned that this fossil is old. Indeed it is. It dates back roughly 133 –140 million years, which places it in the lower Cretaceous. What is remarkable is that it predates the previous record holder by something like 11 million years. Even more remarkable, however, is what this tiny fossil can tell us about the ecology of Pinus at that time. 

Firstly, the leaf scars indicate that this tree had two needles per fascicle. This implies that the genus Pinus had already undergone quite the adaptive radiation by this time. If this is the case, it pushes back the clock on pine evolution even earlier. Another interesting feature are the presence of resin ducts. In extant species, these ducts secrete highly flammable terpenes, which would have potentially promoted fire. 

Species that exhibit this morphology today often utilize an ecology that promotes devastating crown fires that clear the land of competition for their seedlings. Although more evidence is needed to confirm this, it nonetheless suggests that such fire adaptations in pines were already shaping the landscape of the Cretaceous period. All in all, this fossil is a reminder that big things often come in small packages. 

Photo Credit: Howard Falcon-Lang, Royal Holloway University of London

Further Reading:

http://bit.ly/1QP85zm

Aquatic Angiosperm: A Cretaceous Origin?

Via Bernard Gomeza, Véronique Daviero-Gomeza, Clément Coiffardb, Carles Martín-Closasc, David L. Dilcherd, and O. Sanisidro [SOURCE]

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.

Via Bernard Gomeza, Véronique Daviero-Gomeza, Clément Coiffardb, Carles Martín-Closasc, David L. Dilcherd, and O. Sanisidro [SOURCE]

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

Carboniferous Remains

My recent fossil hunting excursion has found me a bit obsessed with the paleobotanical world. As a child, I cut my teeth on biology through the lens of fossilized remains of ancient sea creatures. However, this trip turned up something quite special for a plant lover like myself. Among seams of coal we were finding a bewildering amount of fossilized plant life. However, these were no ordinary plants. What we were finding were the remains of some of the earliest forests on our planet. The rocks we were searching in date back to the Carboniferous Era, over 300 million years before the present. For most of the day I was quite excited by what looked like fossilized fern fronds. It was only after I got home and did some research that I realized how special these fossils really are. 

Illustration by Styracosaurus Rider

Though they resemble the ferns we find throughout the world today, these were not ferns as we know them. What we had found eternalized by mineralization were the remains of a group of plants that no longer exist. These, my friends, were the seed ferns. This lineage arose during the Devonian but really hit their peak abundance during the Carboniferous and Permian eras. Evolutionarily speaking, the seed ferns sit somewhere between true ferns and cycads on the tree of life. The clue to their position here are the presence of seeds, something ferns don't have but cycads do. The seed ferns are only distantly related to ferns and more closely related to cycads. Many believe they represent the earliest gymnosperms. 

Hanging off of the branches are what are referred to as "cupules."Inside the cupules sat the embryos. It is believed that these were wind pollinated as most fossilized cupules show an opening at one end where pollen would have entered. As it stands currently, the seed ferns are the first plants known to reproduce via seeds and therefore represent a major step in the evolution of plants as a whole. Seeds meant that unlike ferns, which require moist conditions to reproduce, the seed ferns were able to colonize a wider swath of habitat types, especially those that were much drier than the swamps in which terrestrial plants evolved.

Why the seed ferns went extinct while both ferns and cycads didn't is not entirely clear. It is likely that competition with more derived seed bearing plants certainly didn't help matters. Despite their disappearance, the legacy of seed ferns lives on today in the fossil fuels we are so hopelessly addicted to and the fossil evidence they leave behind. At the very least, pondering over their fossilized remains is something everyone should take the time to do. 

Photo Credit: [1] [2]

Further Reading: [1] [2] [3] [4]