Apocynaceae Ant House

Bullate leaves help the vine clasp to the tree as well as house ant colonies. Photo by Richard Parker licensed under CC BY-NC-SA 2.0

The dogbane family, Apocynaceae, comes in many shapes, sizes, and lifestyles. From the open-field milkweeds we are most familiar with here in North America to the cactus-like Stapeliads of South Africa, it would seem that there is no end to the adaptive abilities of this family. Being an avid gardener both indoors and out, the diversity of Apocynaceae means that I can be surrounded by these plants year round. My endless quest to grow new and interesting houseplants was how I first came to know a genus within the family that I find quite fascinating. Today I would like to briefly introduce you to the Dischidia vines.

The genus Dischidia is native to tropical regions of China. Like its sister genus Hoya, these plants grow as epiphytic vines throughout the canopy of warm, humid forests. Though they are known quite well among those who enjoy collecting horticultural curiosities, Dischidia as a whole is relatively understudied. These odd vines do not attach themselves to trees via spines, adhesive pads, or tendrils. Instead, they utilize their imbricated leaves to grasp the bark of the trunks and branches they live upon.

The odd, bulb-like leaves of the urn vine (Dischidia rafflesiana) Photo by Bernard DUPONT licensed under CC BY-SA 2.0

One thing we do know about this genus is that most species specialize in growing out of arboreal ant nests. Ant gardens, as they are referred to, offer a nutrient rich substrate for a variety of epiphytic plants around the world. What's more, the ants will visciously defend their nests and thus any plants growing within.

The flowers of Dischidia ovata Photo by Krzysztof Ziarnek, Kenraiz licensed under CC BY-SA 4.0

Some species of Dischidia take this relationship with ants to another level. A handful of species including D. rafflesiana, D. complex, D. major, and D. vidalii produce what are called "bullate leaves." These leaves start out like any other leaf but after a while the edges stop growing. This causes the middle of the leaf to swell up like a blister. The edges then curl over and form a hollow chamber with a small entrance hole.

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

These leaves are ant domatia and ant colonies quickly set up shop within the chambers. This provides ample defense for the plant but the relationship goes a little deeper. The plants produce a series of roots that crisscross the inside of the leaf chamber. As ant detritus builds up inside, the roots begin to extract nutrients. This is highly beneficial for an epiphytic plant as nutrients are often in short supply up in the canopy. In effect, the ants are paying rent in return for a place to live.

Growing these plants can take some time but the payoff is worth. They are fascinating to observe and certainly offer quite a conversation piece as guests marvel at their strange form.

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

Further Reading: [1]

The Extraordinary Catasetum Orchids

Male Catasetum osculatum. Photo by Orchi licensed under CC BY-SA 3.0

Orchids, in general, have perfect flowers in that they contain both male and female organs. However, in a family this large, exceptions to the rules are always around the corner. Take, for instance, orchids in the genus Catasetum. With something like 166 described species, this genus is interesting in that individual plants produce either male or female flowers. What's more, the floral morphology of the individual sexes are so distinctly different from one another that some were originally described as distinct species. 

Female Catasetum osculatum. Photo by Valdison Aparecido Gil licensed under CC BY-SA 4.0

In fact, it was Charles Darwin himself that first worked out that plants of the different sexes were indeed the same species. The genus Catasetum enthralled Darwin and he was able to procure many specimens from his friends for study. Resolving the distinct floral morphology wasn't his only contribution to our understanding of these orchids, he also described their unique pollination mechanism. The details of this process are so bizarre that Darwin was actually ridiculed by some scientists of the time. Yet again, Darwin was right. 

Catasetum longifolium. Photo by Maarten Sepp licensed under CC BY-SA 4.0

If having individual male and female plants wasn't strange enough for these orchids, the mechanism by which pollination is achieved is quite explosive... literally. 

Catasetum orchids are pollinated by large Euglossine bees. Attracted to the male flowers by their alluring scent, the bees land on the lip and begin to probe the flower. Above the lip sits two hair-like structures. When a bee contacts these hairs, a structure containing sacs of pollen called a pollinia is launched downwards towards the bee. A sticky pad at the base ensures that once it hits the bee, it sticks tight. 

Male Catasetum flower in action. Taken from BBC's Kingdom of Plants.

Bees soon learn that the male flowers are rather unpleasant places to visit so they set off in search of a meal that doesn't pummel them. This is quite possibly why the flowers of the individual sexes look so different from one another. As the bees visit the female flowers, the pollen sacs on their back slip into a perfect groove and thus pollination is achieved. 

The uniqueness of this reproductive strategy has earned the Catasetum orchids a place in the spotlight among botanists and horticulturists alike. It begs the question, how is sex determined in these orchids? Is it genetic or are there certain environmental factors that push the plant in either direction? As it turns out, light availability may be one of the most important cues for sex determination in Catasetum. 

Photo by faatura licensed under CC BY-NC-ND 2.0

A paper published back in 1991 found that there were interesting patterns of sex ratios for at least one species of Catasetum. Female plants were found more often in younger forests whereas the ratios approached an even 1:1 in older forests. What the researchers found was that plants are more likely to produce female flowers under open canopies and male flowers under closed canopies. In this instance, younger forests are more open than older, more mature forests, which may explain the patterns they found in the wild. It is possible that, because seed production is such a costly endeavor for plants, individuals with access to more light are better suited for female status. 

Catasetum macrocarpum. Photo by maarten sepp licensed under CC BY-SA 2.0

Aside from their odd reproductive habits, the ecology of these plants is also quite fascinating. Found throughout the New World tropics, Catasetum orchids live as epiphytes on the limbs and trunks of trees. Living in the canopy like this can be stressful and these orchids have evolved accordingly. For starters, they are deciduous. Most of the habitats in which they occur experience a dry season. As the rains fade, the plants will drop their leaves, leaving behind a dense cluster of green pseudobulbs. These bulbous structures serve as energy and water stores that will fuel growth as soon as the rains return. 

Catasetum silvestre in situ. Photo by Antonio Garces licensed under CC BY-NC-ND 2.0

The canopy can also be low in vital nutrients like nitrogen and phosphorus. As is true for all orchids, Catasetum rely on an intimate partnership with special mychorrizal fungi to supplement these ingredients. Such partnerships are vital for germination and growth. However, the fungi that they partner with feed on dead wood, which is low in nitrogen. This has led to yet another intricate and highly specialized relationship for at least some members of this orchid genus. 

Photo by faatura licensed under CC BY-NC-ND 2.0

Mature Catasetum are often found growing right out of arboreal ant nests. Those that aren't will often house entire ant colonies inside their hollowed out pseudobulbs. This will sometimes even happen in a greenhouse setting, much to the chagrin of many orchid growers. The partnership with ants is twofold. In setting up shop within the orchid or around its roots, the ants provide the plant with a vital source of nitrogen in the form of feces and other waste products. At the same time, the ants will viciously attack anything that may threaten their nest. In doing so, they keep many potential herbivores at bay.  

Female Catasetum planiceps. Photo by sunoochi licensed under CC BY 2.0

To look upon a flowering Catasetum is quite remarkable. They truly are marvels of evolution and living proof that there seems to be no end to what orchids have done in the name of survival. Luckily for most of us, one doesn't have to travel to the jungles and scale a tree just to see one of these orchids up close. Their success in the horticultural trade means that most botanical gardens house at least a species or two. If and when you do encounter a Catasetum, do yourself a favor and take time to admire it in all of its glory. You will be happy that you did. 

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9] 

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

Live-In Mites

Photo by Scott Zona licensed under CC BY-NC 2.0

Hearing the word "mite" as a gardener instantly makes me think of pests such as spider mites. This is not fair. The family to which mites belong (Acari) is highly varied and contains many beneficial species. Many mites are important predators at the micro scale. Some are fungivorous, eating potentially harmful species of fungi. Whereas this may be lost on the majority of us humans, it is certainly not lost on many species of plants. In fact, the relationship between some plants and mites is so strong that these plants go as far as to provide them with a sort of home.

Photo by Jsarratt licensed under CC BY-SA 3.0

Domatia are specialized structures that are produced by plants to house arthropods. A lot of different plant species produce domatia but not all of them are readily apparent to us. For instance, many trees and vines such as red oak (Quercus rubra), sugar maples (Acer saccharum), black cherries (Prunus serotina), and many species of grape (Vitis spp.) produce tiny domatia specifically for mites. The domatia are often small, hairy, and function as shelter for both the mites and their eggs.

By housing certain species of mites, these plants are ensuring that they have a steady supply of hunters and cleaners living on their leaves. Predatory mites are voracious hunters, keeping valuable leaves free of microscopic herbivores while frugivorous mites clean the leaves of detrimental fungi that are known to cause infections such as powdery mildew. The exchange is pretty straight forward. Mites get a home and a place to breed and the plants get some protection. Still, some plants seem to want to sweeten the relationship in a literal sense.

Some plants, specifically grape vines in the genus Vitis, also produce extrafloral nectaries on their leaves. These tiny glands secrete sugary nectar. In a paper recently published in the Annals of Botany, it was found that extrafloral nectaries enhances the efficacy of these mite domatia by enticing more mites to stick around. By adding nectar to domatia-producing leaves that did not secrete it, the researchers found that nectar increases beneficial mite densities on these leaves by 60 - 80%. This translates to an increase in fitness for these plants in the long run.

I love research like this. I had no idea that so many of my favorite and most familiar tree and vine species had entered into an evolutionary relationship with beneficial mites. This adds a whole new layer of complexity to the interactions within any given environment. It just goes to show you how much is left to be discovered in our own back yards.

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

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

Orchid Ant Farms

Photo by Scott Zona licensed under CC BY-NC 2.0

I am beginning to think that there is no strategy for survival that is off-limits to the orchid family. Yes, as you may have figured out by now, I am a bit obsessed with these plants. Can you really blame me though? Take for instance Schomburgkia tibicinis (though you may also see it listed under the genera Laelia or more accurately, Myrmecophila). These North, Central, and South American orchids are more commonly referred to as cow-horn orchids because they possess hollow pseudobulbs that have been said to been used by children as toy horns. What is the point of these hollow pseudobulbs?

A paper published back in 1989 in the American Journal of Botany found the answer to that question. As it turns out, ants are quite closely associated with orchids in this genus. They crawl all over the flowers, feeding on nectar. The relationship goes much deeper though. If you were to cut open one of these hollow pseudobulbs, you would find ant colonies living within them. The ants nest inside and often pile up great stores of food and eventually waste within these chambers. The walls of the chambers are lined with a dark tissue that was suspect to researchers.

Using radioactively labeled ants, the researchers found that the orchids were actually taking up nutrients from the ant middens! What's more, nutrients weren't found solely in adjacent tissues but also far away, in the actively growing parts of the roots. These orchids are not only absorbing nutrients from the ants but also translocating it to growing tissues.

While orchids without a resident ant colony seem to do okay, it is believed that orchids with a resident ant colony do ever so slightly better. This makes sense. These orchids grow as epiphytes on trees, a niche that is not high in nutrients. Any additional sources of nutrients these plants can get will undoubtedly aid in their long-term survival. Also, because the ants use the orchids as a food source and a nest site, they are likely defending them from herbivores.

Photo Credit: Scott.Zona (http://bit.ly/1hvWiGX)

Further Reading:
http://www.jstor.org/stable/2444355