The Largest Mistletoe

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When we think of mistletoes, we generally think about those epiphytic parasites living on branches way up in the canopy. The mistletoe we are discussing in this post, however, is a decent sized tree. Nuytsia floribunda is a native of western Australia where it is known locally as moojar or the Christmas tree. To the best of our knowledge, it is the largest mistletoe known to science.

Nuytsia floribunda is a member of the so-called showy mistletoe family (Loranthaceae). It along with all of its mistletoe cousins reside in the order Santalales but from a phylogenetic standpoint, the family Loranthaceae is considered sister to all other mistletoes. This has excited my botanists as it allows us a chance to better understand how parasitism may have evolved in this group as a whole.

Speaking of parasitism, there are some incredible things going on with N. floribunda that are worth talking about. For starters, it is not fully parasitic but rather hemiparasitic. As you can tell by looking at the tree decked out in a full canopy of leaves, N. floribunda is entirely capable of photosynthesizing on its own. In fact, experts feel that it is fully capable of meeting all of its own carbohydrate needs. Instead, it parasitizes other plants in order to acquire water and minerals. How it manages this is remarkable to say the least.

Nuytsia floribunda is a root parasite. Its own roots fan out into the surrounding soil looking for other roots to parasitize. Amazingly, exploratory roots of individual N. floribunda have been found upwards of 110 meters (360 ft.) or more away from the tree. When N. floribunda do find a suitable host root, something incredible happens. It begins to form specialized roots called “haustoria”, which to form a collar-like structure around the host’s roots.

Whole haustoria of Nuytsia (white [ha]) and host root (dark brown). * indicates `gland' and developing `cutting device.

Whole haustoria of Nuytsia (white [ha]) and host root (dark brown). * indicates `gland' and developing `cutting device.

The collar gradually swells and a small horn forms on the inside of the haustoria. Swelling of the haustoria is the result of an influx of water and as the pressure around the host root builds, the haustorial horn of N. floribunda physically cuts into its victim. Once this cut is formed, the haustoria form balloon-like outgrowths which intrude into the xylem tissues of the host root, thus forming the connection that allows N. floribunda to start stealing the water and minerals it needs.

Even more amazing is the fact that roots aren’t the only thing that N. floribunda will attempt to exploit. Many inanimate objects have been found wrapped up in a haustorial embrace including dead twigs, rocks, fertilizer granuals, and even electric cables! Its non-selective parasitic nature appears to have left it open to exploring other, albeit dead end options. I don’t want to paint the picture that this tree as the enemy of surrounding vegetation. It is worth noting that N. floribunda extracts very little from any given host so its impact is spread out among the surrounding vegetation, making its overall impact on host plants minimal most of the time.

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Provided its needs have been met, N. floribunda puts on one heck of a show around December. In fact, the timing of its blooms is the reason it earned the common name of Christmas tree. Flowering for this species is not a modest affair. Each tree is capable of producing multiple meter-long inflorescences decked out in sprays of bright orange to yellow flowers. The flowers themselves produce copious amounts of pollen and nectar, making it an important food source for resident pollinators. Though many different species have been documented visiting the flowers, it is thought that beetles and wasps are the most effective at pollination.

Seed dispersal for N. floribunda is mainly via wind. Each fruit is adorned with three prominent wings. After they detach from the tree, the fruits usually break apart into three samaras, each with its own wing. The key for success of any propagule is ending up in a site suitable for germination. According to some, this can be a bit tricky and attempts at cultivating this plant in captivity have not been terribly successful. It would seem that nature knows best when it comes to reproductive success in N. floribunda. It may be worth trying to figure it out though because recent evidence suggests that this species is not faring well with human development. As the surrounding landscapes of western Australia become more and more urbanized, plants like N. floribunda seem to be on the decline. Perhaps renewed interest in growing this species could change the tide for it as well as others.

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

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

The Desert Mistletoe: Evolution In Action

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There are a multitude of mistletoes on this planet (for example: 1, 2, 3) and all of them are parasites to one degree or another. I find parasitic plants absolutely fascinating because there are as many variations on this lifestyle as there are hosts to parasitize. On a recent botanical adventure in the Sonoran Desert, I met yet another representative of this group - the desert mistletoe (Phoradendron californicum). Once I knew what I was looking at, I could not wait to do some research. As it turns out, this species has garnered quite a bit of attention over the years and it is teaching us some interesting tidbits on how parasites may evolve.

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The desert mistletoe is not hard to spot, especially during the driest parts of the year when most of its host trees have shed their leaves. It looks like a leafless, tangled mass of pendulous stems sitting among the branches of larger shrubs and trees. It can be found growing throughout both the Mojave and Sonoran deserts and appears to prefer leguminous trees including palo verde (Parkinsonia florida), mesquite (Prosopis spp.), and Acacia.

The desert mistletoe is a type of hemiparasite, which means it is capable of performing photosynthesis but nonetheless relies on its host tree for water and other nutrients. Lacking leaves, the desert mistletoe meets all of its photosynthetic needs via its green stems. Its leafless habit also makes its flowers and fruit all the more conspicuous. Despite their small size, its flowers are really worth closer inspection. When in bloom, a desert mistletoe comes alive with the hum of various insects looking for energy-rich nectar and pollen. Even before you spot them, you can easily tell if there is a blooming mistletoe nearby as the flowers give off a wonderfully sweet aroma. It appears that the desert mistletoe takes no chances when it comes to reproduction in such an arid climate.

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As I mentioned above, the desert mistletoe has been the subject of inquiry over the last few decades. Researchers interested in how parasitic plants evolve have illuminated some intriguing aspects of the biology of this species, especially as its relates to host preference. It would appear that our interest in this species seems to be situated at an important time in its evolutionary history. Not all populations of desert mistletoe "behave" in the same way. In fact, each seems to be heading towards more intense specialization based on its preferred host.

By performing seed transplant experiments, researchers have demonstrated that various populations of desert mistletoe seem to be specializing on specific tree species. For instance, when seeds collected from mistletoe growing on Acacia were placed on paleo verde or mesquite, they experienced significantly less germination than if they were placed on another Acacia. Though the exact mechanisms aren't clear at this point in time, evidence suggests that the success of desert mistletoe may be influenced by various hormone levels within the host tree, with isolated populations becoming more specialized on the chemistry of their specific host in that region.

Speaking of isolation, there is also evidence to show that populations of desert mistletoe growing on different host trees are reproductively isolated as well. Populations growing on mesquite trees flower significantly later than populations growing on Acacia or palo verde. Essentially this means that their genes never have the chance to mix, thus increasing the differences between these populations. Again, it is not entirely certain how the host tree may be influencing mistletoe flowering time, however, hormones and water availability are thought to play a role.

Another intriguing idea, and one that has yet to be tested, are the roles that seed dispersers may play in this evolutionary drama. After pollination, the desert mistletoe produces copious amounts of bright red berries that birds find irresistible. Two birds in particular, the northern mockingbird and the Phainopepla, aggressively defend fruiting mistletoe shrubs within their territories. It could be possible that these birds may be influencing which trees the seeds of the desert mistletoe end up on. Again, this is just a hypothesis but one that certainly deserves more attention.

A Phainopepla on the lookout for mistletoe berries.

A Phainopepla on the lookout for mistletoe berries.

Love them or hate them, there is something worth admiring about mistletoes. At the very least, they are important components of their native ecology. What's more, species like the desert mistletoe have a lot to teach us about the way in which species interact and what that means for biodiversity.

Photo Credit: [1]

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

The Explosive Dwarf Mistletoes

I used to think mistletoes were largely a southern phenomenon, preferring regions with mild or even no winters. Then I was introduced to the dwarf mistletoes in the genus Arceuthobium. These odd parasites can be found growing throughout the northern hemisphere. Their affinity for conifers has landed them on the watch list of many a forester yet, despite their economic implications, the dwarf mistletoes are fascinating parasitic plants. 

First and foremost, these are aggressive little plants. They vary in their host specificity. Some species can grow on a wide variety of conifer species from Abies balsamea (balsam fir), Larix laricina (American larch), to Pinus strobus (eastern white pine), whereas others are more specialized, preferring only spruces (Picea spp.). Regardless, infestations of these parasites can do some interesting things to conifer stands. 

Similar to other mistletoes, the dwarfs are stem parasites. They penetrate into their hosts vascular tissues and set up shop, sucking up water and photosynthates and giving nothing in return. Because of this, large infestations can seriously drain their host trees as they themselves have reduced or even no photosynthetic capacity. Additionally, they interfere with nutrient and hormone flows throughout the branches of their host. Such disruptions can result in the formation of dense clusters of branches called "witches brooms." Some dwarf mistletoe infestations can become so intense that they effectively girdle their host tree.

In natural settings, this serves an ecological function. By weakening their hosts, dwarf mistletoes can leave room for other plant species to take root. They also keep one species from becoming too dominant. As such, mistletoe infestations can actually increase plant diversity in the long run. Dwarf mistletoe infestations only become an issue once humans get involved. They can cause serious financial issues for foresters as well as damage important or valued specimen trees. In our highly fragmented forests, their natural behavior can get in the way of human ideals. 

All of this talk of damage can distract us from just how amazing some of these species really are from an organismal standpoint. For instance, the lodgepole pine dwarf mistletoe, Arceuthobium americanum, is capable of thermogenesis. Unlike the other examples of thermogenesis in the plant world, this has nothing to do with flowers. Instead, thermogenesis in A. americanum is used as a seed dispersal agent. 

The dwarf mistletoes don't rely on fleshy fruits to get their seeds from one tree to another. Instead, they utilize ballistic means. As their seed pods mature, they gradually swell. Once pressure is great enough, the seed pods erupt, sending their sticky seeds flying through the canopy at speeds of up to 62 mph (100 km/h)! If lucky, the seeds will stick to the branches of a viable host or be transported there in the fur or feathers of an animal. For A. americanum, the eruption of its seed pods is triggered by heat. Using specialized metabolic pathways at the cellular level, A. americanum is able to heat its seed pods up to ~2 °C warmer than its surroundings, thus triggering its pods to explode. 

Pretty incredible for a species so often labelled as a pest. 

Photo Credit: [1]

Further Reading: [1] [2] [3] [4]

The Holoparasitic Mistletoes

Flowers of   Tristerix aphyllu s

Flowers of Tristerix aphyllus

The order collectively referred to as mistletoes is incredibly diverse. They range in size from rather large trees down to little more than a couple leaves, barely recognizable on their hosts. Even more unique are the mistletoes that have foregone much of what we would readily recognize as an actual plant. These parasitic plants have adopted an endophytic lifecycle, living their entire lives within the vascular tissues of their host plants, only visible to observers when in flower. 

Tristerix aphyllus is one such species. Its hosts are cacti in the genus Echinopsis (formerly Trichocereus) native to Columbia and Chile. Being an endophyte, the majority of this mistletoe lives as a mycelial-like network of filaments that wrap around the vascular tissues of the host cactus. The only part of the mistletoe that ever emerges are the flowers. They come in both red and yellow forms. What may appear to be lovely cactus covered in red flowers are actually the flowers of Tristerix. Strangely enough, the occasional small leaf is produced on the flowering branches. Though there is chlorophyll in the leaf, researchers believe that they perform little if any photosynthesis.

Fruiting   Tristerix aphyllu s

Fruiting Tristerix aphyllus

This is not a parasitic relationship that is unique to cacti either. Africa has its own endoparasitic mistletoe as well. However, as we have discussed before, Africa does not have any native cacti (http://on.fb.me/1zPbac7). Instead, through convergent evolution, plants in the genus Euphorbia have followed similar adaptive trajectories. As such, at least one species of African mistletoe has followed suit.

Flowers of   Viscum minimum

Flowers of Viscum minimum

A species known scientifically as Viscum minimum finds the cactus-like Euphorbia horrida and E. polygona to its liking. Like Tristerix, Viscum minimum is endoparasitic, living entirely within the tissues of its Euphorbia host until it decides to flower. It too produces brightly colored berries that aid in its dispersal to a new host. 

The main seed dispersers are birds. After consumption, a bird either regurgitates the embryo or passes it out the other end. If that bird happens to be sitting on a host cactus or Euphorbia, the embryo will grow into a seedling that quickly taps into its new host and begins its internal parasitic life. It will not be seen again until it flowers.

Viscum minimum  beginning to set seed.

Viscum minimum beginning to set seed.

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


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