Gymnosperms and Fleshy "Fruits"

 Fleshy red aril surrounding the seeds of  Taxus baccata.

Fleshy red aril surrounding the seeds of Taxus baccata.

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.

 Cross section of a  Ginkgo  ovule with red arrow showing the integument.

Cross section of a Ginkgo ovule with red arrow showing the integument.

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)

“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.

Ginkgo “fruits” are full of butyric acid and smell like rotting butter or vomit.

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 ).

“Fruits” contained within the female cone of a cycad (Lepidozamia peroffskyana).

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 .

Berry-like seeds of Gnetum gnemon.

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 .

Red, fleshy bracts of Ephedra distachya.

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 .

The berry-like cones of Juniperus communis.

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

Cedrus deodara

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.

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

Cedrus libani

  Cedrus libani

Cedrus libani

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

Male cones of Cedrus atlantica

 Female  Cedrus  cones.

Female Cedrus cones.

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.

  Cedrus atlantica

Cedrus atlantica

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

Picea_pungens_USDA3.jpg

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 and has since followed its own evolutionary trajectory.

 Male cones are short-lived but quite attractive.

Male cones are short-lived but quite attractive.

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. 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.

The typical female cone of the Colorado blue spruce.

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. Not all trees produce the same amount or consistency of wax and therefore not all look 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 and Norway maples, 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. Though one could certainly do worse than planting this conifer, it nonetheless overshadows more ecologically friendly tree choices. 

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

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