Krassilovia: An Amazing Cretaceous Conifer

Reconstructing extinct organisms based on fossils is no simple task. Rarely do paleontologists find complete specimens. More often, reconstructions are based on fragments of individuals found either near one another or at least in similar rock formations. This is especially true for plants as their growth habits frequently result in fragmentary fossilization. As such, fossilized plant remains of a single species are often described as distinct species until subsequent detective work pieces together a more complete picture.

Such was the case for the fossil remains of what were described as Krassilovia mongolica and Podozamites harrisii. Hailing from the Early Cretaceous (some 100-120 million years ago), Krassilovia was only known from oddly spiny cone scales and Podozamites was only known from strap-shaped leaves found in a remote region of Mongolia. Little evidence existed to suggest they belonged to the same plant. That is, until these structures were analyzed using scanning electron micrographs.

(A–C) Articulated seed cones, (D) Isolated cone axis, (E) Incomplete leafy shoot showing a cluster of three attached leaves, (F) Three detached strap-shaped leaves, G) Detail of A showing tightly imbricate interlocking bract-scale complexes, (H) Detail of leaf apex showing converging veins, (I) Three isolated bract-scale complexes showing abaxial (top) and adaxial (bottom) surfaces, (J) Two isolated seeds showing narrow wings. [SOURCE]

These fossilized plant remains were preserved in such detail that microscopic anatomical features such as stomata were visible under magnification. By studying the remains of these plants as well as others, scientists discovered some amazing similarities in the stomata of Krassilovia and Podozamites. Unlike other plant remains associated with those formations, the Krassilovia cone scales and Podozamites leaves shared the exact same stomate morphology. Though not without some uncertainty, the odds that these two associated structures would share this unique morphological trait by chance is slim and suggests that these are indeed parts of the same plant.

The amazing discoveries do not end with stomata either. After countless hours of searching, fully articulated Krassilovia cones were eventually discovered, which finally put the strange spiky cone scales into context. It turns out those spiked scales interlocked with one another, with the two bottom spikes of one scale interlocking with the three top spikes of the scale below it. In life, such interlocking may have helped protect the developing seeds within until they had matured enough to be released. Also, the sheer volume of cone scales coupled with other minute anatomical details I won’t go into here indicate that, similar to Abies and Cedrus cones, Krassilovia cones completely fell apart when fully ripe.

Though not related, the cone scales of the extinct Krassilovia (left) show similarities with the cone scales of modern day Cryptomeria species (right).

Interestingly, the ability to resolve microscopic structures in these fossils has also provided insights into some modern day taxonomic confusion. It turns out that Krassilovia shares many minute anatomical similarities with present day Gnetales. Gnetales really challenge our perception of gymnosperms and their superficial resemblance to angiosperms have led many to suggest that they represent a clade that is sister to flowering plants. However, more recent molecular work has placed the extant members of Gnetales as sister to the pines. Evidence of shared morphological features between extinct conifers like Krassilovia and modern day Gnetales add some interesting support to this hypothesis. Until more concrete evidence is described and analyzed, the true evolutionary relationships among these groups will remain the object of heated debate for the foreseeable fture.

What we can say is that Krassilovia mongolica was one remarkable conifer. Its unique morphology clearly demonstrates that conifers were once far more diverse in form and function than they are currently. Even the habitat in which Krassilovia once lived is not the kind of place you can find thriving conifer communities today. Krassilovia once grew in a swampy habitat. However, whereas only a few extant conifers enjoy swamps, Krassilovia once shared its habitat with a wide variety of conifer species, the likes of which we are only just beginning to appreciate. I for one am extremely excited to see what new fossil discoveries will uncover in the future.

LISTEN TO EPISODE 300 OF THE IN DEFENSE OF PLANTS PODCAST TO LEARN MORE ABOUT THIS FOSSIL AND THE ECOSYSTEM IN WHICH IT ONCE EXISTED.

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

Further Reading: [1]

Fraser Fir: A New Look at an Old Friend

Photo by James St. John licensed under CC BY 2.0

Growing up, Fraser fir (Abies fraseri) was a fairly common sight in our house. Each winter this species would usually win out over other options as the preferred tree for our living room during the holiday season. Indeed, its pleasing shape, lovely color, and soft needles have made it one of the most popular Christmas trees around the world. Amazingly, despite its popularity as a decoration, Fraser fir is so rare in the wild that it is considered an endangered species.

Fraser fir is native to only a handful of areas in the southern Appalachian Mountains. Together with red spruce (Picea rubens), this conifer makes up one of the rarest ecosystems on the continent - the southern Appalachian spruce-fir forest. Such forests only exist at elevations above 4,000 ft (1,200 m) from southwestern Virginia to western North Carolina and eastern Tennessee. The reason for this limited distribution is rooted in both modern day climate and North America’s glacial past.

USGS/Public Domain

Whereas anyone hiking through Appalachian spruce-fir forests could readily draw similarities to boreal forests found farther north, the Appalachian spruce-fir forests are nonetheless unique, hosting many species found nowhere else in the world. Indeed, these forests are holdovers from the Pleistocene when the southeast was much cooler than it is today. As glaciers retreated and the climate warmed, Appalachian spruce-fir forests “retreated” up the mountains, following their preferred climate zones until they hit the peaks of mountains and couldn’t go any further.

Indeed, Fraser fir is in large part limited in its distribution by temperature. This conifer does not perform well at high temperatures and is readily out-competed by other species under warmer conditions. Another factor that has maintained Appalachian spruce-fir forests at elevation is fog. The southern Appalachian Mountains host eastern North America’s only temperate rainforest and fog commonly blankets high elevation areas throughout the year. Research has shown that in addition to keeping these areas cool, fog also serves as an important source of water for Fraser fir and its neighbors. As fog condenses on its needles, these trees are able to absorb that water, keeping them hydrated even when rain is absent.

A view of an Appalachian spruce-fir forest from the Blue Ridge Parkway.

Due to its restricted habitat, Fraser fir has never been extremely common. However, things got even worse as Europeans colonized North America. Over the past two centuries, unsustainable logging and grazing practices have decimated southern Appalachian spruce-fir forests, fragmenting them into even smaller patches with no connectivity in between. In areas where thin, rocky soils were not completely washed away, Fraser fir seedlings did return, however, this was not always the case. In areas where soils were were lost, southern Appalachian spruce–fir forests were incapable of regenerating.

If the story ended there, Fraser fir and its habitat would still be in trouble but sadly, things only got worse with the introduction of the invasive balsam woolly adelgid (Adelges piceae) from Europe around 1900. Like the hemlock woolly adlegid, this invasive, sap-feeding insect has decimated Fraser fir populations throughout southern Appalachia. Having shared no evolutionary history with the adelgid, Fraser fir is essentially defenseless and estimates suggest that upwards of 90% of infect trees have been killed by the invasion. Although plenty of Fraser fir seedlings have sprung up in the wake of this destruction, experts fear that as soon as those trees grow large enough to start forming fissures in their bark, the balsam woolly adelgid will once again experience a massive population boom and repeat the process of destruction again.

Dead Fraser fir as seen from Clingman’s Dome. Photo by Brian Stansberry licensed under CC BY 3.0

The loss of Fraser fir from this imperiled ecosystem has had a ripple effect. Fraser fir is much sturdier than its red spruce neighbors and thus provides an important windbreak, protecting other trees from the powerful gusts that sweep over the mountain tops on a regular basis. With a decline in the Fraser fir canopy, red spruce and other trees are more susceptible to blowdowns. Also, the dense, evergreen canopy of these Appalachian spruce-fir forests produces a unique microclimate that fosters the growth of myriad mosses, liverworts, ferns, and herbs that in turn support species like the endangered endemic spruce-fir moss spider (Microhexura montivaga). As Fraser fir is lost from these areas, the species that it once supported decline as well, placing the whole ecosystem at risk of collapse.

The moss-dominated understory of an Appalachian spruce-fir forest supports species found nowhere else in the world. Photo by Miguel.v licensed under CC BY 3.0

Luckily, the plight of this tree and the habitat it supports has not gone unnoticed by conservationists. Numerous groups and agencies are working on conserving and restoring Fraser fir and southern Appalachian spruce-fir forests to at least a portion of their former glory. This is not an easy task by any means. Aside from lack of funding and human power, southern Appalachian spruce-fir forest conservation and restoration is hindered by the ever present threat of a changing climate. Fears that the life-giving fog that supports this ecosystem may be changing make it difficult to prioritize areas suitable for reforestation. Also, the continued threat from invasive species like the balsam woolly adelgid can hamper even the best restoration and conservation efforts. Still, this doesn’t mean we must give up hope. With continued collaboration and effort, we can still ensure that this unique ecosystem has a chance to persist.

Please visit the Central Appalachian Spruce Restoration Initiative (CASRI) website to learn more!

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

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

Maxipiñon: One of the Rarest Pines in the World

Photo by Ruff tuff cream puff licensed under public domain

The maxipiñon (Pinus maximartinezii) is one of the rarest pines on Earth. A native of southern Sierra Madre Occidental, Mexico, nearly all individuals of this species can be found scattered over an area that collectively spans only about 3 to 6 square miles (5 – 10 km²) in size. Needless to say, the maxipiñon teeters on the brink of extinction. As a result, a lot of effort has been put forward to better understand this species and to develop plans aimed at ensuring it is not lost forever.

The maxipiñon has only been known to science for a few decades. It was described back in 1964 after botanist Jerzy Rzedowski noted some exceptionally large pine seeds for sale at a local market. He named the species in honor of Maximino Martínez, who contributed greatly to our understanding of Mexican conifers. However, it was very obvious that the maxipiñon was well known among the residents of Zacatecas.

The reason for this are its seeds. The maxipiñon is said to produce the largest and most nutritious seeds of all the pines. As such, it is a staple of the regional diet. Conversations with local farmers suggest that it was much more common as recent as 60 years ago. Since then, its numbers have been greatly reduced. It soon became apparent that in order to save this species, we had to learn a lot more about what threatens its survival.

The most obvious place to start was recruitment. If any species is to survive, reproduction must outpace death. A survey of local markets revealed that a lot of maxipiñon seeds were being harvest from the wild. This would be fine if maxipiñon were widespread but this is not the case. Over-harvesting of seeds could spell disaster for a species with such small population sizes.

Indeed, surveys of wild maxipiñon revealed there to be only 2,000 to 2,500 mature individuals and almost no seedlings. However, mature trees do produce a considerable amount of cones. Therefore, the conclusion was made that seed harvesting may be the single largest threat to this tree. Subsequent research has suggested that seed harvests actually may not be the cause of its rarity. It turns out, maxipiñon population growth appears to be rather insensitive to the number of seeds produced each year. Instead, juvenile tree survival seems to form the biggest bottleneck to population growth.

Photo by Krzysztof Ziarnek, Kenraiz licensed under CC BY-SA 4.0

You see, this tree appears to be more limited by suitable germination sites than it does seed numbers. It doesn’t matter if thousands of seeds are produced if very few of them ever find a good spot to grow. Because of this, scientists feel that there are other more serious threats to the maxipiñon than seed harvesting. However, humans are still not off the hook. Other human activities proved to be far more damaging.

About 50 years ago, big changes were made to local farming practices. More and more land was being cleared for cattle grazing. Much of that clearing was done by purposefully setting fires. The bark of the maxipiñon is very thin, which makes it highly susceptible to fire. As fires burn through its habitat, many trees are killed. Those that survive must then contend with relentless overgrazing by cattle. If that wasn’t enough, the cleared land also becomes highly eroded, thus further reducing its suitability for maxipiñon regeneration. Taken together, these are the biggest threats to the ongoing survival of this pine. Its highly fragmented habitat no longer offers suitable sites for seedling growth and survival.

As with any species this rare, issues of genetic diversity also come into play. Though molecular analyses have shown that maxipiñon does not currently suffer from inbreeding, it has revealed some interesting data that give us hints into the deeper history of this species. Written in maxipiñon DNA is evidence of an extreme population bottleneck that occurred somewhere between 400 and 1000 years ago. It appears that this is not the first time this tree has undergone population decline.

There are a few ways in which these data can be interpreted. One is that the maxipiñon evolved relatively recently from a small number of unique and isolated individuals. Perhaps a hybridization event occurred between two closely related piñon species - the weeping piñon (Pinus pinceana) and Nelson piñon (Pinus nelsonii). Another possibility, which does not rule out hybridization, is that the maxipiñon may actually be the result of artificial selection by agriculturists of the region. Considering the value of its seeds today, it is not hard to imagine farmers selecting and breeding piñon for larger seeds. It goes without saying that these claims are largely unsubstantiated and would require much more evidence to say with any certainty, however, there is plenty of evidence that civilizations like the Mayans were conserving and propagation useful tree species much earlier than this.

Despite all we have learned about the maxipiñon over the last few decades, the fate of this tree is far from secure. Ex situ conservation efforts are well underway and you can now see maxipiñon specimens growing in arboreta and botanical gardens around the world. Seeds from these populations are being used for storage and to propagate more trees. Sadly, until something is done to protect the habitat on which it relies, there is no telling how long this species will last in the wild. This is why habitat conservation efforts are so important. Please support local land conservation efforts in your area because the maxipiñon is but one species facing the loss of its habitat.

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

Further Reading [1] [2] [3]

Gymnosperms and Fleshy "Fruits"

Fleshy red aril surrounding the seeds of Taxus baccata. Photo by Frank Vincentz licensed under the GNU Free Documentation License.

Many of us were taught in school that one of the key distinguishing features between gymnosperms and angiosperms is the production of fruit. Fruit, by definition, is a structure formed from the ovary of a flowering plant. Gymnosperms, on the other hand, do not enclose their ovules in ovaries. Instead, their unfertilized ovules are exposed (to one degree or another) to the environment. The word “gymnosperm” reflects this as it is Greek for “naked seed.” However, as is the case with all things biological, there are exceptions to nearly every rule. There are gymnosperms on this planet that produce structures that function quite similar to fruits.

Internal anatomy of a Ginkgo ovule with red arrow showing the integument.

Photo copyright Bruce Kirchoff, Licensed under CC BY 2.0

The key to understanding this evolutionary convergence lies in understanding the benefits of fruits in the first place. Fruits are all about packing seeds into structures that appeal to the palates of various types of animals who then eat said fruits. Once consumed, the animals digest the fruity bits and will often deposit the seeds elsewhere in their feces. Propagule dispersal is key to the success of plants as it allows them to not only to complete their reproductive cycle but also conquer new territory in the process. With a basic introduction out of the way, let’s get back to gymnosperms.

“Fruits” of Cephalotaxus fortunei (Cephalotaxaceae)

There are 4 major gymnosperm lineages on this planet - the Ginkgo, cycads, gnetophytes, and conifers. Each one of these groups contains members that produce fleshy structures around their seeds. However, their “fruits” do not all develop in the same way. The most remarkable thing to me is that, from a developmental standpoint, each lineage has evolved its own pathway for “fruit” production.

Ginkgo “fruits” are full of butyric acid and smell like rotting butter or vomit. Photo by H. Zell licensed under CC BY-SA 3.0

For instance, consider ginkgos and cycads. Both of these groups can trace their evolutionary history back to the early Permian, some 270 - 280 million years ago, long before flowering plants came onto the scene. Both surround their developing seed with a layer of protective tissue called the integument. As the seed develops, the integument swells and becomes quite fleshy. In the case of Ginkgo, the integument is rich in a compound called butyric acid, which give them their characteristic rotten butter smell. No one can say for sure who this nasty odor originally evolved to attract but it likely has something to do with seed dispersal. Modern day carnivores seem to be especially fond of Ginkgo “fruits,” which would suggest that some bygone carnivore may have been the main seed disperser for these trees.

“Fruits” contained within the female cone of a cycad (Lepidozamia peroffskyana). Photo by Tony Rodd licensed under CC BY-NC-SA 2.0

The Gnetophytes are represented by three extant lineages (Gnetaceae, Welwitschiaceae, and Ephedraceae), but only two of them - Gnetaceae and Ephedraceae - produce fruit-like structures. As if the overall appearance of the various Gnetum species didn’t make you question your assumptions of what a gymnosperm should look like, its seeds certainly will. They are downright berry-like!

Berry-like seeds of Gnetum gnemon. Photo by gbohne licensed under CC BY-SA 2.0

The formation of the fruit-like structure surrounding each seed can be traced back to tiny bracts at the base of the ovule. After fertilization, these bracts grow up and around the seed and swell to become red and fleshy. As you can imagine, Gnetum “fruits” are a real hit with animals. In the case of some Ephedra, the “fruit” is also derived from much larger bracts that surround the ovule. These bracts are more leaf-like at the start than those of their Gnetum cousins but their development and function is much the same.

Red, fleshy bracts of Ephedra distachya. Photo by Le.Loup.Gris licensed under CC BY-SA 3.0

Whereas we usually think of woody cones when we think of conifers, there are many species within this lineage that also have converged on fleshy structures surrounding their seeds. Probably the most famous and widely recognized example of this can be seen in the yews (Taxus spp.). Ovules are presented singly and each is subtended by a small stalk called a peduncle. Once fertilized, a group of cells on the peduncle begin to grow and differentiate. They gradually swell and engulf the seed, forming a bright red, fleshy structure called an “aril.” Arils are magnificent seed dispersal devices as birds absolutely relish them. The seed within is quite toxic so it usually escapes the process unharmed and with any luck is deposited far away from the parent plant.

The berry-like cones of Juniperus communis. Photo by Piero Amorati, ICCroce - Casalecchio di Reno, Bugwood.org licensed under Creative Commons Attribution 3.0 License.

Another great example of fleshy conifer “fruits” can be seen in the junipers (Juniperus spp.). Unlike the other gymnosperms mentioned here, the junipers do produce cones. However, unlike pine cones, the scales of juniper cones do not open to release the seeds inside. Instead, they swell shut and each scale becomes quite fleshy. Juniper cones aren’t red like we have seen in other lineages but they certainly garnish the attention of many a small animal looking for food.

I have only begun to scratch the surface of the fruit-like structures in gymnosperms. There is plenty of literary fodder out there for those of you who love to read about developmental biology and evolution. It is a fascinating world to uncover. More importantly, I think the fleshy “fruits” of the various gymnosperm lineages stand as a testament to the power of natural selection as a driving force for evolution on our planet. It is amazing that such distantly related plants have converged on similar seed dispersal mechanisms by so many different means.

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8]

Further Reading: [1] [2] [3] [4] [5] [6] [7]

The Only True Cedars

Cedrus deodara. Photo by PabloEvans licensed under CC BY 2.0

The only true cedars on this planet can be found growing throughout mountainous regions of the western Himalayas and Mediterranean. All others are cedars by name only. The so-called “cedars” we encounter here in North America are not even in the same family as the true cedars. Instead, they belong to the Cypress family (Cupressaceae). The true cedars all belong to the genus Cedrus and are members of the family Pinaceae. They are remarkable trees with tons of ecological and cultural value.

J. White,1803-1824.

The true cedars are stunning trees to say the least. They regularly reach heights of 100 ft. (30 m.) or more and can live for thousands of years. Cedars exhibit a dimorphic branching structure, with long shoots forming branches and smaller shoots carrying most of the needle load. The needles themselves are borne in dense, spiral clusters, giving the branches a rather aesthetic appearance. Each needle produces layers of wax, which vary in thickness depending on how exposed the tree is growing. This waxy layer helps protect the tree from sunburn and desiccation.

Cedrus libani. Photo by Zeynel Cebeci licensed under CC BY-SA 4.0

Cedrus libani. Photo by Leonid Mamchenkov licensed under CC BY 2.0

One of the easiest ways to identify a cedar is by checking out its cones. All members of the genus Cedrus produce upright, barrel-shaped cones. Male cones are smaller and don’t stay on the tree for very long. Female cones, on the other hand, are quite large and stay on the tree until the seeds are ripe. Upon ripening, the entire female cone disintegrates, releasing the seeds held within. Each seed comes complete with blisters full of distasteful resin, which is thought to deter seed predators.

Male cones of Cedrus atlantica. Photo by Meneerke bloem licensed under CC BY-SA 3.0

Female Cedrus cones. Photo by Zeynel Cebeci licensed under CC BY-SA 4.0

In total, there are only four recognized species of cedar - the Atlas cedar (Cedrus atlantica), the Cyprus cedar (C. brevifolia), the deodar cedar (C. deodara), and the Lebanon cedar (C. libani). I have heard arguments that C. brevifolia is no more than a variant of C. libani but I have yet to come across any source that can say this for certain. Much more work is needed to assess the genetic structure of these species. Even their place within Pinaceae is up for debate. Historically it seems that Cedrus has been allied with the firs (genus Abies), however, work done in the early 2000’s suggests that Cedrus may actually be sister to all other Pinaceae. We need more data before anything can be said with certainty.

Regardless, two of these cedars - C. atlantica & C. libani - are threatened with extinction. Centuries of over-harvesting, over-grazing, and unsustainable fire regimes have taken their toll on wild populations. Much of what remains is not considered old growth. Gone is the heyday of giant cedar forests. Luckily, many populations are now located in protected areas and reforestation efforts are being put into place throughout their range. Still, the ever present threat of climate change is causing massive pest outbreaks in these forests. The future for these trees hangs in the balance.

Photo Credit: Wikimedia Commons

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

I've Got the Colorado Blues

Dave Powell, USDA Forest Service (retired), Bugwood.org licensed under a Creative Commons Attribution 3.0 License.

You would be hard pressed to find a resident of temperate North America who has never seen a Colorado blue spruce. These iconic trees are a staple of every sapling give-away and can be found in countless landscape plans all over the continent. There is no denying the fact that the blue hues of Picea pungens have managed to tap into the human psyche and in doing so has managed to spread far beyond its relatively limited range. However, despite its popularity, few people ever really get to know this species. Even fewer will ever encounter it in the wild. Today I would like to introduce you to a brief natural history of Picea pungens. 

Despite its common name, P. pungens is not solely a denizen of Colorado. It can be found in narrow swaths of the Rocky Mountains of Wyoming, Idaho, south to Utah, northern and eastern Arizona, southern New Mexico, and of course, central Colorado. There are also some rumored populations in Montana as well. It has a very narrow range compared to its more common relative, the Engelmann spruce (Picea engelmannii). Whereas some authors consider the Colorado blue spruce to be a subspecies of the Engelmann spruce, the paucity of natural hybrids where these two species overlap suggests otherwise. It is likely that Colorado blue spruce split off from this lineage at some point in the past and has been following its own evolutionary trajectory ever since.

Female cones are quite attractive when they emerge. Photo by JJ Harrison (https://www.jjharrison.com.au) licensed under CC BY-SA 3.0

One of the reasons P. pungens has become such a popular landscape tree is due to its extreme hardiness. Indeed, this is one sturdy tree species. Not only can it handle drought, P. pungens is also capable of surviving temperatures as low as -40 degrees Celsius with minimal foliar damage. Little stands in the way of a well established Colorado blue. In the wild it can be found growing on gentle mountain slopes at elevations of 6,000 to 10,000 feet (1,800 to 3000 m). It is also a long lived and highly fecund tree. The most highly productive seed years for P. pungens begin at age 50 and last until it reaches roughly 150 years of age. Seeds germinate best on bare soils, which probably keeps this species limited to these mountainous areas in the wild.

The typical female cone of the Colorado blue spruce. Photo by U.S. Fish and Wildlife Service Public Domain

Another component of its landscape popularity is its characteristic blue color. In reality, not all trees exhibit this coloration. Its blue hue is the result of epicuticular wax deposits on the leaves as they are produced in the spring. Individual trees rpduce varying amounts and consistencies of wax and therefore may not appear blue. Wax production seems to be controlled by a genetic factor and therefore is often a shared trait among isolated populations. The wax functions as sun screen, reflecting harmful UV rays away from sensitive developing foliage. This is why it is most prominent in new growth. The wax can and often does degrade over the span of a growing season, resulting in duller trees come fall. 

Despite how interesting this spruce is, Picea pungens, in my opinion, represents the epitome of lazy landscaping. Like Norway spruce (Picea abies) and Norway maples (Acer platanoides), P. pungens seems to be an all-too-easy choice for those looking to save a quick buck. As a result, countless numbers of these trees line streets and demarcate property boundaries. Though P. pungens is native to North America, its narrow home range makes its ecological function elsewhere quite minimal. Sure, one could certainly do worse than planting this conifer, but it nonetheless overshadows more ecologically friendly tree choices. If you are looking to add a new tree to your landscape, take a few minutes to search for more ecologically friendly species that are native to your region.

Photo Credit: [1] [2] [3]

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