A Herbaceous Conifer From the Triassic

aleth1.jpg

It is hard to make broad generalizations about groups of related organisms. There are always exceptions to any rule. Still, there are some “facts” we can throw around that seem to apply pretty well to specific branches on the tree of life. For instance, all of the gymnosperm lineages we share our planet with today are woody, relatively slow to reach sexual maturity, and are generally long-lived. This has not always been the case. Fossil discoveries from France suggest that in the past, gymnosperms were experimenting with a more herbaceous lifestyle.

The fossils in question were discovered in eastern France back in the 1800’s. The strata from which they were excavated dates back to the Middle Triassic, some 247 million years ago. Immortalized in these rocks were numerous spindly plants with strap-like leaves and a few branches, each ending in what look like tiny cones. Early interpretations suggested that these may represent an extinct lycopod, however, further investigation suggested something very surprising - a conifer with an herbaceous growth habit.

Indeed, thanks to even more scrutiny, it is now largely agreed upon that what was preserved in these rocks were essentially herbaceous conifers. The fossils were given the name Aethophyllum stipulare. They are wonderfully complete, depicting roots, shoots, leaves, and reproductive organs. Moreover, the way in which they were fossilized preserved lots of fine-scale anatomical details. Taken together, there are plenty of clues available that allow paleobotanists to say a lot about how this odd conifer made a living.

For starters, they were not very big plants. Not a single specimen has been found that exceeds 2 meters (6.5 ft) in height. The main stem of these conifers only seem to branch a couple of times. Cones were formed at the tips of the upper branches and not a single specimen has been found that depicts subsequent growth following cone formation. This suggests that Aethophyllum exhibited determinate growth, meaning that individuals grew to a certain size, reproduced, and did not continue to grow after that. Female cones were situated at the tips of the upper most branches and male cones were situated at the tips of lower shoots. The smallest reproductive individuals that have been unearthed are only 30 cm (11 in) in height, which suggests that Aethophyllum  was capable of reproducing within a few months of germination.

Artists reconstruction of  Aethophyllum stipulare

Artists reconstruction of Aethophyllum stipulare

Amazingly, researchers were also able to extract fossilized pollen and seeds from some of the Aethophyllum cones. The pollen itself is saccate, much like what we see in many extant conifers. By comparing the morphology of the pollen extracted from the cones to other fossil pollen records, researchers now feel confident that Aethophyllum is the source of pollen grains discovered in sediments from western, central, and southern Europe, Russia, Northern Africa, and China, suggesting that Aethophyllum was pretty wide spread during the Middle Triassic. Aethophyllum seeds were small, ellipsoid, and were not winged, likely germinating a short distance from the parent.

The stems of Aethophyllum are interesting in the own right. Thanks to their preservation, cross sections have been made and they reveal that these plants only ever produced secondary tracheids and primary xylem. The only place on the plant where any signs of woody secondary xylem occur are at the base of the cones. This adds further confirmation that Aethophyllum was herbaceous at the onset of sexual maturity.

Another intriguing aspect of the stem is the presence of numerous large air spaces within the stem pith. Today, this anatomical feature is present in plants like bamboo, Equisetum, and the flowering stalks of Agave, all of which exhibit alarmingly fast growth rates for plants. This suggests that not only did Aethophyllum reproduce early in its life, it also likely grew extremely fast.

1. Smallest fertile plant in the Grauvogel and Gall collections, with two stems extending from the root, and terminal ovulate cone (OC) on one branch (scale bar=10 cm). 2. Cross-section of stem in the Grauvogel and Gall collections showing cauline bundles with scanty wood (at left, top and right) surrounding large pith with large, aerenchymatous lacunae and interspersed pith parenchyma cells. Vascular cambium, phloem, and more peripheral tissues are not preserved (scale bar=200 μm). 3.Seedling in the Grauvogel and Gall collections showing primary root (R), cotyledons (C) and stem (S) with apically borne leaves (scale bar=10 cm).  Quoted from SOURCE

1. Smallest fertile plant in the Grauvogel and Gall collections, with two stems extending from the root, and terminal ovulate cone (OC) on one branch (scale bar=10 cm). 2. Cross-section of stem in the Grauvogel and Gall collections showing cauline bundles with scanty wood (at left, top and right) surrounding large pith with large, aerenchymatous lacunae and interspersed pith parenchyma cells. Vascular cambium, phloem, and more peripheral tissues are not preserved (scale bar=200 μm). 3.Seedling in the Grauvogel and Gall collections showing primary root (R), cotyledons (C) and stem (S) with apically borne leaves (scale bar=10 cm). Quoted from SOURCE

Mature Aethophyllum aren’t the only fossils available either. Many seedlings have been discovered in close proximity to the adults. Seedlings were also exquisitely preserved, depicting hypocotyl, a primary root system, two two-veined cotyledons, and a short stem with four-veined leaves arranged in a helix. The fact that seedlings and adults were found in such close proximity lends to the idea that Aethophyllum populations were made up of multi-aged stands, not unlike some of the early successional plants we find in disturbed habitats today.

The sediments in which these plants were fossilized can also tell us something about the habitats in which Aethophyllum grew. The rock layers are made up of a mix of sediments typical of what one would find in a flood plain or delta. Also, Aethophyllum aren’t the only plant remains discovered. Many species known to grow in regularly disturbed, flood-prone habitats have also been found. Taken together these lines of evidence suggest that Aethophyllum was similar to what we would expect from herbaceous plants growing in similar habitats today. They grew fast, reproduced early, and had to jam as many generations in before the next flood ripped through and hit the reset button.

Aethophyllums small size, lack of wood, and rapid growth rate all point to a ruderal lifestyle. Today, this niche is largely filled by angiosperms. No conifers alive today can claim such territories. The discovery of Aethophyllum demonstrates that this was not always the case. The fact that pollen has been found far outside of France suggests that this ruderal lifestyle worked quite well for Aethophyllum.

The terrestrial habitats of the Middle Triassic were dominated by the distant relatives of modern day ferns, lycophytes, and gymnosperms. Needless to say, it was a very different world than anything that we are familiar with today. However, that does not mean that the pressures of natural selection were necessarily different. Aethophyllum is evidence that specific selection pressures, in this case regular flood disturbance, select for similar traits in plants through time. Why Aethophyllum went extinct is anyone’s guess. Despite how well they have been preserved, there is still a lot of mystery surrounding this plant.

Photo Credit: [1]

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



Fossilized Flower Places Angiosperms in the Jurassic

1, style branches; 2, dendroid style; 3, sepal; 4, ovarian roof; 5, scale; 6, seed; 7, cup-form receptacle/ovary; 8, bract; 9, petal; 10, unknown organ (staminode?).  [SOURCE]

1, style branches; 2, dendroid style; 3, sepal; 4, ovarian roof; 5, scale; 6, seed; 7, cup-form receptacle/ovary; 8, bract; 9, petal; 10, unknown organ (staminode?). [SOURCE]

Despite their dominance on the landscape today, the origin of flowering plants is shrouded in mystery. The odds of any living material becoming fossilized is extremely rare and when you consider the delicate and ephemeral nature of most flowers, one can begin to understand why their fossils are so special. The last few decades have seen tantalizing evidence emerge from fossil beds dating to the Cretaceous Period but a recent set of fossils from China predate the oldest confirmed angiosperm fossils by 50 million years. That’s right, it would appear that flowering plants were already on the scene by the early Jurassic!

The fossils in question have been coined Nanjinganthus dendrostyla. They were discovered in China in a formation that dates back roughly 174 million years. To most of us they look like a bunch of dark, albeit elaborate smudges on the rocks. To a trained eye, however, these smudges reveal intricate anatomical details. Amazingly, the team of paleobotanists responsible for this discovery had a lot of material to work with. Descriptions were made on a whopping 264 specimens representing 198 individual flowers. This amount of data means that the declaration of angiosperm affinity stands on pretty solid ground.

A single  Nanjinganthus  flower  [SOURCE]

A single Nanjinganthus flower [SOURCE]

Aside from their age, there is a lot about these fossils that surprised researchers. Probably the biggest surprise is their overall appearance. Paleobotanists have long hypothesized that early angiosperm flowers likely resembled something akin to a modern day Magnolia and invoke floral features such as apocarpy, a superior ovary, and a lack of an obvious style as likely features to look for in ancient plant fossils. Surprisingly, Nanjinganthus does not seem to conform to many of these expectations.

One of the most striking features of these fossils are the styles. They are large and branched like tiny trees (hence the specific epithet “dendrostyla”). The tree-like appearance of the style suggests that early angiosperms likely did not rely on insects for pollination. The branches themselves greatly increase the amount of surface area available for pollen capture, which could mean that Nanjinganthus was wind pollinated.

Flowers of  Nanjinganthus  preserved in different states and their details. For specific details on each image, please see   SOURCE

Flowers of Nanjinganthus preserved in different states and their details. For specific details on each image, please see SOURCE

Another surprising feature is the presence of an inferior ovary that, by its very definition, sits below the sepals and petals. It has long been hypothesized that early angiosperms would exhibit superior ovaries so this discovery means that we must rethink our expectations of how flowers evolved. For instance, it suggests we may not be able to make broad inferences on the past based on what we see in extant angiosperm lineages. It could also suggest that the origin of flowering plants was not a single event but rather a series of individual occurrences. It could also be the case that the origin of flowering plants occurred much earlier than the Jurassic and that Nanjinganthus represents one of many derived forms. Only further study and more fossils can help us answer such questions.

Another way in which Nanjinganthus deviates from theoretical expectations is in the presence of both sepals and petals. Up until now, paleobotanists have been fond of the idea that petals arose much later in angiosperms, having evolved over time as leaves became more and more specialized for attracting pollinators. The fact that Nanjinganthus was likely wind pollinated yet had both sepals and petals is a bit of a conundrum and further emphasizes the need to revisit some of our long-held assumptions of flowering plant evolution.

Details of the sepal and petal as seen through different forms of microscopic analysis. For specific details on each image, please see  SOURCE .

Details of the sepal and petal as seen through different forms of microscopic analysis. For specific details on each image, please see SOURCE.

By far the most important feature present in these fossils are the ovaries. For any fossil to unequivocally qualify as an angiosperm, it must have seeds encased in an ovary. This, after all, is the main feature that separates angiosperms from gymnosperms. Indeed, Nanjinganthus does appear to fit this definition. Thanks to the sheer amount of fossils available for study, the team discovered that the seeds of Nanjinganthus were enclosed in a cup-like chamber that was sealed off from the outside world by a structure they refer to as an “ovarian roof.” This roof does not appear to have any sort of opening, which worked out quite nicely for paleobotanists as it prevented sediments from entering into the chamber, thus preserving the seeds or ovules (it is hard to tell where they were in the developmental process) for study. This feature more than all others secures its identity as a flowering plant.

Based on the sediments in which these flowers were fossilized, it appears that this plant grew close to water. Also, despite its abundance in this particular fossil layer, it very likely was not a common component of this Jurassic landscape. In reality we still have a lot to learn about Nanjinganthus. What we can say with some certainty at this point is that the presence of Nanjinganthus in the early Jurassic likely means that flowering plants arose even earlier. Nanjinganthus is most definitely not the first flower. We will probably never find the first of anything. It is an ancient flower though, predating all other discoveries by at least 50 million years. This is why paleontology is so incredible. Who knows what the next blow of a rock hammer will turn up!

 

EDIT (10/27/2018): Since writing this post it has come to my attention that there is quite a bit of controversy attached to the description of this fossil. Many have reached out informing me that these fossils may actually be a gymnosperm organ rather than a flower. Despite all of the outcry I have yet to see any published critiques on this particular controversy. I anxiously await more professional input on the subject but for now I have decided to keep the content of the original piece as is. Of course extraordinary claims require extraordinary evidence and not being a paleobotanist myself, I cannot trust hearsay on the internet as fact, no matter how vociferous, until I see it published in a peer reviewed outlet of some sort. Please stay tuned as this story develops! 

Photo Credits: [1]

Further Reading: [1]

The Rise and Fall of the Scale Trees

SS2139210.jpg

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

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 the started branching, young trees would have resembled a bushy, green bottle brush.

Juvenile scale tree on the left & the adult on the right

Juvenile scale tree on the left & the adult on the right

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

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.

leps.PNG

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

The fossilized strobilus of a Lepidodendron

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”

A fossilized root or “stigmaria”

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] [3] [4] [5] [6] [7]

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