Meet the She-Oaks

628814411_ed018d2c14_o.jpg

No, what you are looking at here is not a type of conifer. Nor is it an oak. This odd plant belongs in its own family - Casuarinaceae. Despite their gymnosperm appearance, this is in fact a family of flowering plants. Though the name “she-oak” does hint at their larger position within the order Fagales, it was actually given to these trees in reference to the density of their wood in comparison to more commonly harvested oak species. Other common names for trees in this group include ironwood, bull-oak, beefwood.

As a whole this family sorts out as sister to Myricaceae in the order Fagales. It' is comprised of 4 genera (Allocasuarina, Casuarina, Ceuthostoma, and Gymnostoma) and roughly 91 species spread among Australia, Malaysia, and much of Polynesia. It is extremely difficult to make generalizations across so many species but there is one aspect of this family that makes them stand out - their appearance.

Gymnostoma  sp.

Gymnostoma sp.

Gymnostoma nobile  in Sarawak, Malaysia.

Gymnostoma nobile in Sarawak, Malaysia.

Without close inspection, one could be forgiven for thinking the various Casuarinaceae were species of conifer. For starters, their leaves have been reduced to tiny whorls surrounding their photosynthetic stems. The stems themselves have taken up the role of photosynthetic organs, which is one of the reasons this family is known for its drought tolerance. Reducing the surface area available for gas exchange helps to reduce water loss in the process. The stems themselves are arranged with whorls around the branches, giving them a rather bunched appearance. The photosynthetic branches are sometimes referred to as being ‘equisetiform’ as they superficially resemble the stems of Equisetum. They do not shed their photosynthetic branches and are therefore evergreen.

As mentioned, these are flowering plants. Their flowers themselves are aggregated into spike-like inflorescences near the tips of branches. Clusters of male flowers resemble catkin-like strobili and are often brightly colored. Female flowers are clustered into a more ovoid shape, with long, filamentous pistils sticking out like fiery, red pompoms. After fertilization, bracts at the base of the female flowers swell and the whole inflorescence starts to look more like some sort of a conifer cone than anything floral. This may have to do with the fact that, like conifers, the various Casuarinaceae are wind pollinated. Therefore, their reproductive structures have had to deal with similar selective forces related to optimizing pollen dispersal and capture.

Casuarina equisetifolia  with catkin-like male flowers and smaller, red female flowers.

Casuarina equisetifolia with catkin-like male flowers and smaller, red female flowers.

Allocasuarina distyla  female flowers and infructescence.

Allocasuarina distyla female flowers and infructescence.

Another interesting trait common to Casuarinaceae is the ability to fix nitrogen. The plants themselves don’t do the fixing, rather they form specialized nodules on their roots that house nitrogen-fixing bacteria. Unlike perennial legumes that regrow their nodules year after year, the members of Casuarinaceae hold onto their nodules, which can grow into impressive structures over time. This ability to house nitrogen-fixing bacteria is also shared with other members of the order Fagales, including members of Betulaceae and Myricaceae.

Thanks to the fact that they can tolerate drought, fix nitrogen, and have high timber value, species of Casuarinaceae have been introduced far outside of their native ranges. This has created yet another invasive species issue. Various Casuarinaceae have become serious pests in places like Central and South America, the Carribbean, and the Middle East. Control of such hardy plants can be extremely difficult once they reach a critical mass that maintains them on the landscape. Keep you eye out for these species. Not only are they interesting in their own right, knowing them can help you better understand their role in ecosystems both native and not.

Allocasuarina decaisneana  (Desert Oaks), Central Australia

Allocasuarina decaisneana (Desert Oaks), Central Australia

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

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

The Early Days Of A Symbiosis?

aglao10.jpg

Despite the ubiquitous nature of symbioses across the globe, evidence of their origins is scant to say the least. Mostly we look for clues of their origin hidden within the fossil record. Excitingly, a series of fossils discovered in Scotland reveal what very well be the early days of plant-cyanobacterial interactions. Thanks to these exquisitely preserved fossils, we now have the earliest record of an association between these two groups of organisms.

The fossils themselves date back to the early Devonian, some 400 million years ago. They hail from a hot spring community which allowed wonderfully detailed preservation of everything down to the cellular level. Needless to say, this was a drastically different time for life on this planet. Plants were really starting to dominate the landscape. In the case of the fossil discoveries in question, one plant in particular is the star of this show. 

Meet Aglaophyton major. This odd looking plant would have been a common site in these sorts of habitats. It largely consisted of a small, leafless stem that branched as it ambled over the ground. These stems bore the stomata, which allowed gas exchange to occur. Every once in a while, a stem would throw up a reproductive structure called a sporangium, which housed the spores. At the ground level, the stems would occasionally produce root-like rhizoids that have been found in association with fossilized mycorrhizal fungi in the soil.

In total, A. major only stood about 18 cm in height. Though abundant, it was relatively small compared to some of the other vegetation coming online at this point in time. It is likely that A. major could tolerate occasional flooding. In fact, some have speculated that flooding may have been necessary for the germination of its spores. It's this periodic inundation with water that likely led to an interesting and tantalizing relationship with cyanobacteria. 

1. Transverse section through two typical axes showing the simple internal organization; slide P1828; bar = 1 mm. 2. Anatomy of the prostrate mycorrhizal axis (E = epidermis; OC = outer cortex; MAZ = mycorrhizal arbuscule-zone; IC = inner cortex; PIT = phloem-like tissue; CT = conducting tissue); slide P1612; bar = 150 μm. 3. Dense aggregate of cyanobacterial filaments in an area where the axis is injured and has exuded some type of wound secretion (opaque mass); slide P1289; bar = 100 μm. 4. Detail of Plate I, 3, showing part of the cyanobacterial aggregate; bar = 100 μm. 5. Intercellular cyanobacterial filaments near the mycorrhizal arbuscule-zone of the cortex (darker tissue in lower third of image); slide P3652; bar = 50 μm. 6. Group of filaments passing through the intercellular system of the outer cortex; slide P3652; bar = 20 μm.

1. Transverse section through two typical axes showing the simple internal organization; slide P1828; bar = 1 mm. 2. Anatomy of the prostrate mycorrhizal axis (E = epidermis; OC = outer cortex; MAZ = mycorrhizal arbuscule-zone; IC = inner cortex; PIT = phloem-like tissue; CT = conducting tissue); slide P1612; bar = 150 μm. 3. Dense aggregate of cyanobacterial filaments in an area where the axis is injured and has exuded some type of wound secretion (opaque mass); slide P1289; bar = 100 μm. 4. Detail of Plate I, 3, showing part of the cyanobacterial aggregate; bar = 100 μm. 5. Intercellular cyanobacterial filaments near the mycorrhizal arbuscule-zone of the cortex (darker tissue in lower third of image); slide P3652; bar = 50 μm. 6. Group of filaments passing through the intercellular system of the outer cortex; slide P3652; bar = 20 μm.

Cyanobacteria are probably best known for their contribution of oxygen to Earth's early atmosphere. What's more, many also fix nitrogen. That is why the fossil discovery of A. major with cyanobacteria in and around its cells is so exciting. These 400 million year old fossils provide the first evidence of a plant and cyanobacteria in an intimate association.

As mentioned above, the fossilization process was so thorough that it preserved subcellular structures. After thin sectioning some A. major stems, a team of researchers found filaments of cyanobacteria in the process of invading the plant and taking up residence. The cyanobacteria appears to be entering the plant through the stomatal openings along the stem. Once inside, the cyanobacteria show signs of colonazation of substomatal chambers as well as intercellular spaces within the plants tissues.

Although the authors cannot say whether this association was mutualistic or not, it nonetheless represents a model situation detailing how such a symbiotic relationship could have evolved in the first place. Because the cyanobacteria in question here is thought to be aquatic, the only way for it to move into the plant would have been during periodic flooding events. The idea that this could be simply an infection following the death of the plant was considered. However, the non-random distribution of cyanobacteria within A. major cells suggests that this relationship was no accident.

For now, the relationship between A. major and cyanobacteria was likely an "on-again–off-again incidental association" centered around flood events. The fact that A. major was already associated with mycorrhizal fungi at this point in Earth's history certainly suggests that the genetic adaptations necessary for symbiotic relationships were already in place. Though it isn't a smoking gun, these fossils provide the earliest evidence of plants' relationship with cyanobacteria.

Photo Credits: [1]

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