An Intruiguing Relationship Between Ants and Cacti

The extrafloral nectaries of  Pachycereus gatesii  appear as tiny red bumps just below the areole.

The extrafloral nectaries of Pachycereus gatesii appear as tiny red bumps just below the areole.

It’s hard to think of a group of plants that are better defended than cacti. Frequently and often elaborately adorned with vicious spines, these succulents make any animal think twice about trying to take a bite. And yet, for some cacti, spines don’t seem to cut it. A surprising amount of species appear to have taken their defense system to a whole new level by recruiting nature’s most tenacious bodyguards, ants.

Plants frequently have a friend in ants. Spend some time observing ants at work and it’s east to see why. These social insects have numbers and strength on their side. Give ants a reason to be invested in your survival and they will certainly see to it that nothing threatens this partnership. For cacti, this involves the secretion of nectar from specialized tissues called extrafloral nectaries.

Extrafloral nectaries are not unique to cacti. A multitude of plant species produce them, often for similar reasons. Ants love a sugary food source and the more reliable that source becomes, the more adamant an ant colony will be at defending it. The odd thing about cacti is that they don’t seem to have settled on a single type of extrafloral nectary to do the trick. In fact, as many as four different types of extrafloral nectaries have been described in the cactus family.

Ants visiting the extrafloral nectaries covering the developing flowers of  Pilosocereus gounellei .

Ants visiting the extrafloral nectaries covering the developing flowers of Pilosocereus gounellei.

Some cacti secrete nectar from highly modified spines. A great example of this can be seen in genera such as Coryphantha, Cylindropuntia, Echinocactus, Ferocactus, Opuntia, Sclerocactus, and Thelocactus. Such spines are usually short and blunt, hardly resembling spines at all. Other cacti secrete nectar from regular looking spines. This adaptation is odd as there does not seem to be anything special about the anatomy of such spines. Examples of this can be seen in genera such as Brasiliopuntia, Calymmanthium, Harrisia, Opuntia, Pereskiopsis, and Quiabentia. Still others secrete nectar from highly reduced leaves that are found at the base of where the spines originate (the areole). Such leaves have been described in Acanthocereus, Leptocereus, Myrtillocactus, Pachycereus, and Stenocereus. They aren’t easy to recognize as leaves either. Most look like tiny scales. Finally, the fourth type of extrafloral nectary comes in the form of specialized regions of the stem tissue. This has been described in genera such as Armatocereus, Leptocereus, and Pachycereus.

Highly modified spines functioning as extrafloral nectaries in  Ferocactus emoryi.

Highly modified spines functioning as extrafloral nectaries in Ferocactus emoryi.

Seemingly normal spines of  Harrisia pomanensis  secreting nectar.

Seemingly normal spines of Harrisia pomanensis secreting nectar.

Regardless of where they form, their function remains much the same. They secrete a form of nectar which ants find irresistible. The more reliable this food source becomes, the more aggressive ant colonies will be in defending it. This is an especially useful form of defense when it comes to small insect herbivores. Whereas spines deter larger herbivores, they don’t do much to deter organisms that can just slip right through them unharmed. Ants also clean the cacti, potentially removing harmful microbes like fungi and bacteria. Though we are only just beginning to understand the depths of this cactus/ant mutualism, what we have discovered already suggests that the relationship between these types of organisms is far more complex than what I have just outlined above.

For instance, it may not just be sugar that the ants are looking for. In arid desert habitats, water may be the most limiting resource for an ant colony and large, succulent cacti are essentially giant water reservoirs. The key is getting to that water. One study that looked at a species of barrel cactus growing in Arizona called Ferocactus acanthodes found that as spring gives way to summer, the concentration of sugars secreted by the extrafloral nectaries decreases. As a result, the nectar becomes far more watery. Amazingly, ant densities on any given barrel cactus actually increased throughout the summer, despite the fact that the nectar was being watered down. Ants are notoriously prone to desiccation so it stands to reason that water, rather than sugar, is the real prize for colonies hanging out on cacti in such hot desert environments.

The incredible floral display of  Ferocactus wislizeni , a species whose reproductive efforts are affected by the types of ants they attract.

The incredible floral display of Ferocactus wislizeni, a species whose reproductive efforts are affected by the types of ants they attract.

Another interesting observation about the cactus/ant mutualism is that it appears that the identity of the ants truly matters. Though defense is the main benefit to the cactus, research suggests that there is a tipping point in how much such defenses benefit cacti. It has been found that although cacti initially benefit from anti-herbivore and cleaning services, extra aggressive ant species can actually drive off potential pollinators. At least one study has shown that when less aggressive ant species tend cacti, they produce more fruits and those fruits contain significantly more seeds than cacti that have been tended by extremely aggressive ant species. This is especially concerning when we think about the growing issue of invasive ants. As more and more non-native ant species displace native ants, this could really tip the balance for some cactus species.

Despite all of the interesting things we have learned about extrafloral nectaries in the family Cactaceae, there are so many questions yet to be answered. For starters, we still do not know how many different taxa produce them in one form or another. It is likely that closer inspection, especially of rare or poorly understood groups, will reveal that far more cacti produce some type of extrafloral nectary. Also, we know next to nothing about the anatomy of the different types of nectaries. How do they differ from one another and how do some, especially those derived from ordinary spines, actually function? Finally, do these nectaries function year round or is there some sort of seasonal pattern to their development and utility. How does this affect the types of ants they attract and how does that in turn affect the survival and reproduction of these cacti? For such a charismatic group of plants as cacti, we still have to much to learn.

Photo Credits: Thanks to Dr. Jim Mauseth and Dr. John Rebman and Dr. Silvia Rodriguez Machado for use of their photos [1] [2] [3]

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

The Prairie Peninsula

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North American prairies are some of the most endangered habitat types on the planet. Once covering vast swaths of the continent representing the arid rain shadow of the Rocky Mountains, the prairies now occupy only 1% of their former range. We have converted most of that land into agriculture and sub-developments. It may surprise many of you to learn that following the retreat of the Pleistocene glaciers, a subset of prairie ecosystem once stretched a lot further east than one would expect for a prairie. This grassland ecosystem ranged far up into the northeast and even met the Atlantic Coast in parts of New Jersey and Long Island. This was known as the prairie peninsula and today its remnants represent some of the rarest prairie ecosystems that North America has left. 

Ecologists believe that the prairie peninsula owed its existence to an intriguing quirk of the climate at that time. During interglacial periods, eastern North America’s climate was much warmer and drier than it is today. Because of this, prairie grasslands were hypothesized to have migrated east, following the recently exposed terminal moraines that the glaciers left in their wake. Moraine soils tend to be composed of unconsolidated till and are quick to drain water,  which provided perfect conditions for prairies to develop. This prairie peninsula preceded the invasion of trees, which now make up the forests that dominate eastern North America. Today we refer to the remnants of this prairie peninsula as "heaths" or "barrens." Despite their rarity, ecologists have long debated whether such habitats are truly echoes of our glacial past or products of a more recent, cultural clearing of the land.

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A relatively recent paper sheds some light on this debate using some pretty clever detective work. The author used a variety of insects, but mainly focused on spittlebugs, to show that these eastern prairies remnants are, indeed, relics of our glacial past. Apparently, insects like leafhoppers, froghoppers, and spittlebugs are often extremely specialized on specific species of plants, mainly grasses. They feed much like aphids do, by sucking the juices out of the plants vascular tissues. Many of these insects are flightless or at least do not travel great distances from where they were born. If one were to find certain species on the eastern prairies, it would provide strong evidence in support of prairie migration from west to east via the glacial moraines.

The evidence suggests exactly that. Eastern pockets of remnant heaths and barrens do in fact host many of these prairie specialists. What is more interesting is that this research has shown that we can track, with some certainty, the migrational patterns of these ecosystems. As expected, the prairies moved in from the west, during an interglacial period much warmer than now. As they moved across the eastern US, they ran into the Appalachian Mountains, which is a formidable barrier to say the least.

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How did the prairies circumvent this obstacle and end up in pockets along the Atlantic coast? Evidence points to sandy sediments in the numerous valleys along the spine of the Appalachians. Most of these sediments no longer exist due to erosion and out-crowding by forests, causing the current disjunction we see in these rare eastern prairie grasslands. It is amazing to me to think that these pockets of  habitat have existed for centuries, despite all the changes we have laid upon the land. These results are a wonderful example of the uniqueness of these habitats and, now more than ever, show us how much these deserve our attention so that maybe they can persist into the future. It also highlights just how special these ecosystems truly are. They are not something created by the hand of man. Instead, these habitats have survived the test of time. Now they must survive us. Support your local land conservancy today!


Map Credit: [1]

Further Reading: [1]

Meeting One of North America's Rarest Oaks

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A post (and photos) by Robbie Q. Telfer

“Every species is a masterpiece, exquisitely adapted to the particular environment in which it has survived.”

-- E.O. mothereffin Wilson

One of the perks of working at The Morton Arboretum is you get to see cool lectures on tree science for free. At one such program, Dr. Mary Ashley from the University of Illinois at Chicago was sharing her research on oak pollen and how far it can travel to fertilize female flowers (far). She looked at not only trees in the Chicago region, but also oaks off the coast of California and in the Chihuahuan Desert of west Texas, as well as throughout Mexico. That latter oak was a shrubby species called Quercus hinckleyi or Hinckley oak. It is able to spread pollen over far distances as well, despite the fact that there are only 123 individuals known to be left. IUCN lists it as Critically Endangered.

As she was telling us this, it occured to me that I would be in West Texas soon to visit my sister-in-law, so afterwards I approached Dr. Ashley and asked if there was any way I could have the coordinates of Q. hinckleyi so that I could visit it, take a selfie, and luxuriate in the presence of something so rare. I made it clear to her that I understood just how important it was to keep this information a secret, because the last thing this relict needs is to be uprooted by poachers. Which I wish wasn’t a concern, but it is.

Dr. Ashley put me in touch with her colleague Janet Backs who graciously shared the coordinates. I could see the plants from Google maps satellite view. There they were. I probably waved at the computer screen sheepishly.

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As I waited for my time to bask in the majesty of botanical greatness, I consulted my copy of Oaks of North America (1985) by Howard Miller and Samuel Lamb to see what the entry for hinckleyi said.

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Notably, it mentions that “This is another of the oaks with no specific value, except as a curiosity.” More on that later.

After much anticipation, the time was upon us. I decided to drive out to the plants in my rental first thing in the morning after getting to Texas. The Chihuahuan Desert is an astounding place that my Illinoisan eyes weren’t altogether prepared for. It is perhaps the most biodiverse desert in the world, and compared to our prairies, woodlands, and wetlands, it feels like a different planet. Some of the cooler plants I got to see were tree cholla (Cholla sp.), Havard’s century plant (Agave havardiana), Wright’s cliffbrake (Pellaea wrightiana), and little buckthorn (Condalia ericoides). And also a family of introduced aoudads with TWO adorable babies. I also got to see my first javelina (as roadkill) and all kinds of birds new to me.

Tree cholla ( Cholla  sp.)

Tree cholla (Cholla sp.)

Havard’s century plant ( Agave havardiana )

Havard’s century plant (Agave havardiana)

Wright’s cliffbrake ( Pellaea wrightiana )

Wright’s cliffbrake (Pellaea wrightiana)

Little buckthorn ( Condalia ericoides )

Little buckthorn (Condalia ericoides)

Aoudads in the distance.

Aoudads in the distance.

Finally I got to the coordinates - luckily google preloaded the directions on my phone because there was absolutely no cell service where I was. I parked and walked to the plants. And lo, I present to you, Quercus hinckleyi.

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It’s in the white oak family, which I guess means more than just “has round leaves.” These leaves look like holly, and even the shed ones on the ground still had some stabbiness left in them. It’s quite diminutive - certainly compared to any oak I’ve ever seen and even by shrub standards. I’d pinch its cheeks if that wouldn’t make my fingers bleed. After getting the pics I needed and doing the atheist’s version of saying a prayer over it, I floated back to my car like a cartoon cat in love.

The rest of the trip was great and I can’t wait to go back.

Since returning, I have shown several of my non-plant nerd friends the pics of hinckleyi and they seem politely impressed but not, like, actually impressed. This is totally understandable! If your experience with plants is on the order of what looks best in a planting or what tastes best in your tummy, this shrub is not for you. After all “it’s only value is as a curiosity.”

I don’t know about that. I feel like it’s value is greater than that for humans - it’s a window into the North American continent before the climate shifted 10,000 years ago, it’s an individual member of our vast botanical heritage, it is unique, it is adorbs, and it helped Dr. Ashley, and therefore us, understand more things about the movement of oak pollen.

But beyond what it does for US, what if, and hear me out, what if it has a right to existence on its own, without being displaced by pipelines or aoudads or poachers? It is a member of its ecological community, and just like I feel a loss when a member of my community passes, we don’t have the language to articulate what is felt when a member of an ecosystem winks out forever.

Janet Backs told me that she heard of someone who was trying to poach acorns from a subpopulation of hinckleyi and that the landowners where that shrub is actually chased those folks for miles and miles down the road. I love that. I wish every single threatened species/subpopulation had someone who understood its value beyond what it does for humans enough to chase people, possibly with a gun, for miles and miles.

I have had a paltry bucket list for most of my adult life - boring stuff like meeting my heroes or getting to a 7th bowl of never-ending-pasta. But despite their apparent lack of reverence for Q. hinckleyi I think a pretty good guiding list for me would be to visit each of the 77 oaks of North America in their native habitats. I know they won’t all be as special as this experience, but what better way to visit the corners of this continent and its myriad ecological communities, than by visiting each of its oaks? I currently can’t think of any, and would invite anyone to, if not fund me, join me.

An Iris With Multiple Parents

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The Abbeville iris (Iris nelsonii) is a very special plant. It is the rarest of the so-called “Louisiana Irises” and can only be found growing naturally in one small swamp in southern Louisiana. If you are lucky, you can catch it in flower during a few short weeks in spring. The blooms come in a range of colors from reddish-purple to nearly brown, an impressive sight to see siting atop tall, slender stems. However, the most incredible aspect of the biology of this species is its origin. The Abbeville iris is the result of hybridization between not two but three different iris species.

When I found out I would be heading to Louisiana in the spring of 2019, I made sure that seeing the Abbeville iris in person was near the top of my to-do list. How could a botany nut not want to see something so special? Iris nelsonii was only officially described as a species in 1966. Prior to that, many believed hybridization played a role in its origin. Multiple aspects of its anatomy appear intermediate between other native irises. It was not until proper molecular tests were done that the picture became clear.

The Abbeville iris genome contains bits and pieces of three other irises native to Louisiana. The most obvious parent was yet another red-flowering species - the copper iris (Iris fulva). It also contained DNA from the Dixie iris (Iris hexagona) and the zig-zag iris (Iris brevicaulis). If you had a similar childhood as I did, then you may have learned in grade school biology class that hybrids are usually biological dead ends. They may exhibit lots of beneficial traits but, like mules, they are often sterile. Certainly this is frequently the case, especially for hybrid animals, however, more and more we are finding that hybridization has resulted in multiple legitimate speciation events, especially in plants.

How exactly three species of iris managed to “come together” and produce a functional species like I. nelsonii is interesting to ponder. Its three parent species each prefers a different sort of habitat than the others. For instance, the copper iris is most often found in seasonally wet, shady bottomland hardwood forests as well as the occasional roadside ditch, whereas the Dixie iris is said to prefer more open habitats like wet prairies. In a few very specific locations, however, these types of habitats can be found within relatively short distances of eachother.

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Apparently at some point in the past, a few populations swapped pollen and the eventual result was a stable hybrid that would some day be named Iris nelsonii. As mentioned, this is a rare plant. Until it was introduced to other sites to ensure its ongoing existence in the wild, the Abbeville iris was only know to occur in any significant numbers at one single locality. This necessitates the question as to whether or not this “species” is truly unique in its ecology to warrant that status. It could very well be that that single locality just happens to produce a lot of one off hybrids.

In reality, the Abbeville iris does seem to “behave” differently from any of its parental stock. For starters, it seems to perform best in habitats that are intermediate of its parental species. This alone has managed to isolate it enough to keep the Abbeville from being reabsorbed genetically by subsequent back-crossing with its parents. Another mechanism of isolation has to do with its pollinators. The Abbeville iris is intermediate in its floral morphology as well, which means that pollen placement may not readily occur when pollinators visit different iris species in succession. Also, being largely red in coloration, the Abbeville iris receives a lot of attention from hummingbirds.

Although hummingbirds do not appear to show an initial preference when given the option to visit copper and Abbeville irises at a given location, research has found that once hummingbirds visit an Abbeville iris flower, they tend to stick to that species provided enough flowers are available. As such, the Abbeville iris likely gets the bulk of the attention from local hummingbirds while it is in bloom, ensuring that its pollen is being delivered to members of its own species and not any of its progenitors. For all intents and purposes, it would appear that this hybrid iris is behaving much like a true species.

As with any rare plant, its ongoing survival in the wild is always cause for concern. Certainly Louisiana is no stranger to habitat loss and an ever-increasing human population coupled with climate change are ongoing threats to the Abbeville iris. Changes in the natural hydrologic cycle of its swampy habitat appears to have already caused a shift in its distribution. Whereas it historically could be found in abundance in the interior of the swamp, reductions in water levels have seen it move out of the swamp and into ditches where water levels remain a bit more stable year round. Also, if its habitat were to become more fragmented, the reproductive barriers that have maintained this unique species may degrade to the point in which it is absorbed back into an unstable hybrid mix with one or a couple of its parent species. Luckily for the Abbeville, offspring have been planted into at least one other location, which helps to reduce the likelihood of extinction due to a single isolated event.


Photo Credit: [1]

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

The Succulent Passionflowers

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Succulent passionflowers?! It took me a minute to get my head wrapped around the idea. It wasn’t until I saw one in flower that I truly understood. The genus Adenia is found throughout east and west Africa, Southeast Asia, and hits its peak diversity in Madagascar. It comprises approximately 100 species and, as a whole, is poorly understood. Today I would like to introduce you to this bizarre genus within Passifloraceae.

Adenia glauca

Adenia glauca

Adenia is, to date, the second largest genus within the Passionflower family and yet delineating species has been something of a nightmare for botanists over the years. At least some of this confusion lies within the diversity of this odd group. It has been said that few angiosperm lineages surpass Adenia in the diversity of growth forms they exhibit. Though all could be considered succulent to some degree, Adenia runs the gamut from trees to vines, and even tuberous herbs.

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Even within individual species, the overall form of these plants can vary widely depending on the conditions under which they have been growing. Their succulent nature and that fact that many species can reach rather large proportions means that herbarium records for this group are scant at best. Many are only known from a single, incomplete collection of a few bits and pieces of plant. Also, juvenile plants often look very different from their adult forms, making timing of the collection crucial for proper analysis.

To complicate matters more, all Adenia are dioecious, meaning that individual plants are either male or female. Male and female flowers of individual species look pretty distinct and differ a bit from what we have come to expect out of the passionflower family. Often collections were made on only a single sex. This is further complicated by the fact that these plants often exhibit very short flowering seasons. Most come into bloom right before the onset of the rainy season and are entirely leafless at that point in time. Because of this, it has been extremely difficult to accurately match flowering collections to vegetative collections. As such, nearly 1/4 of all Adenia species are missing descriptions of either male or female flowers and their fruits.

Female flower of  Adenia reticulata

Female flower of Adenia reticulata

Male flowers of  Adenia digitata

Male flowers of Adenia digitata

Flowers of  Adenia firingalavensis

Flowers of Adenia firingalavensis

Fruits of  Adenia hondala

Fruits of Adenia hondala

Even genetic work has failed to clear up much of the mysteries that surround this group. Some studies suggest that Adenia is sister to all other genera within Passifloraceae whereas others have even suggested it to be nestled neatly within the genus Passiflora. The most recent work hints at a placement among the tribe Passifloreae. If this confuses you, you are certainly not alone. Until a more complete sampling effort is done on Adenia, I think it is safe to say that this genus will be holding onto its taxonomic mysteries for the foreseeable future.

Adenia globosa

Adenia globosa

All Adenia are perennial plants but how they manage this differs from species to species. Some put all of their energy into underground tubers, producing annual stems and leaves that die back each year. Others don’t produce any tubers and instead store all of their water and nutrients within thick stems. This has made at least a handful of species a hit with succulent growers around the world. It is always an interesting sight to see a giant caudiciform trunk or base with bunches of spindly stems spraying out from the top.

Leaves and fruit of  Adenia cissampeloides

Leaves and fruit of Adenia cissampeloides

Juvenile  Adenia glauca

Juvenile Adenia glauca

Adenia are also extremely toxic plants. The conditions under which these plants evolved are tough and it appears that this group doesn’t want to take any chances on losing any biomass to herbivores. The main class of compounds they produce are called lectins. These proteins cause myriad issues within animal bodies including rapid cell death, blood clotting, inhibition of protein synthesis, and a disruption of ribosome and DNA function. Needless to say, its in any critters best interest to avoid nibbling on any species of Adenia. Even handling and pruning of these plants merits caution.

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Whether you’re a botanist, taxonomist, gardener, or just curious about plant diversity, Adenia is a wonderful example of just how many unknowns are still out there. Regardless of their taxonomic status, these are fascinating species, each with a wonderful ecology and intriguing evolutionary history. These plants are hardy survivors and a great example of the lengths a genus can go to when presented with new opportunities. Undoubtedly many more species await description but the plants we currently know of are fascinating to say the least.

Adenia pechuelii

Adenia pechuelii

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9] [10] [11] [12] [13]

Further Reading: [1] [2]

The Drought Alert System of Terrestrial Plants has an Underwater Origin

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For plants, the transition from water to land was a monumental achievement that changed our world forever. Such a transition was fraught with unique challenges, not the least of which being the ever present threat of desiccation. A new study now suggests that those early land plants already had the the tools to deal with drought and they have their aquatic algal ancestors to thank.

One of the keys to being able to survive drought is being able to detect it in the first place. Without some sort of signalling pathway, plants would not be able to close up stomata and channel vital water and nutrients to more important tissues and organs. As such, elucidating the origins and function of drought signalling pathways in plants has been of great interest to science.

One key set of pathways involved in plant drought response is collectively referred to as the “chloroplast retrograde signaling network.” I’m not even going to pretend that I understand how these pathways operate in any detail but there is one aspect of this network that is the key to this recent discovery. It involves the means by which drought and high-light conditions are sensed in one part of the plant and how that information is then communicated to the rest of the plant. When this signalling pathway is activated, the plant can then begin to produce enzymes that go on to activate defense strategies such as stomatal closure.

Chara braunii  - a modern day example of a streptophyte alga

Chara braunii - a modern day example of a streptophyte alga

The surprise came when researchers at the Australian National University, in collaboration with researchers at the University of Florida, decided to study the chloroplast retrograde signaling network in more detail. They were interested in the inner workings of this process in relation to stomata. Stomata are tiny pores on the leaves and stems of terrestrial plants that regulate the exchange of gases like CO2 and oxygen as well as water vapor. To add some controls to their experiment, the team added a few species of aquatic algae into the mix. Algae do not produce stomata and therefore they reasoned that no traces of chloroplast retrograde signaling network enzymes should be present.

This is not what happened. Instead, the team discovered that the enzymes in question also showed up in a group of algae known as the streptophytes. This was exciting because streptophyte algae hail from the lineage thought to be ancestral to all land plants. It appears that the tools necessary for terrestrial plants to survive drought were already in place before their ancestors ever left the water.

Why this is the case could have something to do with the streptophyte lifestyle. Today, these algae are known to tolerate very tough conditions. Though outright drought is rarely a threat for these aquatic algae, they nonetheless have to deal with scenarios that resemble drought such as high salinity. Streptophyte algae found growing in ephemeral pools must cope with ever increasing concentrations of salinity as the water around them evaporates. It is possible that this drought signalling pathway may have evolved as a response to hyper-saline conditions such as these. Regardless of what was going on during those early days of plant evolution, this research indicates that the ability for terrestrial plants to deal with drought evolved before their ancestors ever left the water.

The closer we look, the more we can appreciate that evolution of important traits isn’t always de novo. More often it appears that new innovations result from a retooling of of older genetic equipment. In the case of land plants, a signalling pathway that allowed their aquatic ancestors to deal with water loss was coopted later on by organs such as leaves and stems to deal with the stresses of life on land. As the old saying goes, “life uhhh… finds a way.”

Photo Credits: [1] [2]

Further Reading: [1] [2]

The Gravel Ghost

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Look closely or you might miss it. The gravel ghost (Atrichoseris platyphylla) is a master of disguise. At home in a small pocket of southwestern North America, this wonderful member of the aster family only puts on a show when rains offer the parched landscape a momentary reprieve.

The gravel ghost is the only member of the genus Atrichoseris. It is different enough from the rest of the chicory tribe (Cichorieae) to warrant its monotypic status. The gravel ghost is a winter annual meaning its seeds germinate at some point in the fall and the plant spends most of the winter putting on growth. As you can probably imagine, life in this corner of the world is pretty tough. Rain is sparse to non-existent and many plants teeter on the edge of desiccation. The fleshy, semi-succulent leaves of the gravel ghost likely store just enough water to offer some insurance against prolonged drought.

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As if drying up wasn’t enough for this plant, the desert’s compliment of hungry herbivores are constantly on the lookout for any plant remotely alive that can offer sustenance. All it takes is a few encounters with the gravel ghost to understand how this plant manages to avoid as much attention as possible. As its common name suggests, this species blends in with the surrounding soil to an extreme degree. From what I can gather, there appears to be a lot of variation in gravel ghost leaf color depending on where the population is growing.

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Some are mostly green whereas others take on a mottled grey hue. Still others seem to have settled on a mixture of browns. It seems that no matter the substrate, the gravel ghost will do its best to blend in. Personally, I would love to see someone investigate what kind of genetic or environmental controls dictate leaf color in this species. It is fascinating to think about how plants can disguise themselves against herbivores.

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Starting in late winter and early spring, the gravel ghost needs to complete its annual life cycle. When rains punctuate the drought, the gravel ghost sends up a spindly inflorescence tipped with a few flower heads. If they are lucky, some stalks will avoid being nipped off by sheep and rabbits. Those that do put on quite a floral display. Each head or ‘capitulum’ explodes with clusters of bright white ray flowers. Only at this point does its affinity with the chicory tribe become apparent.

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The need for such a high impact floral display has everything to do with being an annual. There is only limited time for pollination and seed set. Each gravel ghost must produce enough seeds to enure that at least some survive. They simply don’t have multiple seasons to reproduction. Luckily its a member of the aster family and the opportunity for seed production is usually relatively high. With any luck, plenty of pollinators will find these plants tucked in among rocks and gravel and the process will begin again come that fall.

Photo Credit: Joey (www.instagram.com/crime_pays_but_botany_doesnt)

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



A Hardy Tillandsia That Deserves Our Respect

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As epiphytes go, Tillandsia recuvata (a.k.a. ball moss) doesn’t have the best reputation. All too often it is seen as an unsightly pest of trees that needs to be removed. This could not be farther from the truth. This hardy air plant does no harm to the trees on which it grows. What’s more, its relationship with a specific group of bacteria means it is a major contributor to soil fertility. Today I would like to sing the praise of the indefatigable Tillandsia recuvata.

Tillandsia recuvata is native throughout an impressive chunk of the Americas, from the southern United States through to northern Argentina and Chile. Wherever temperatures rarely dip below freezing, T. recurvata can make an easy living. One of the most remarkable aspects of this species is the array of habitat types in which it grows. This hardy little air plant is equally at home in sub-tropical conditions as it is arid desert habitats. Its ability to tolerate heat, drought, and plenty of air pollution has led to its colonization of urban environments as well.

One of the keys to its success is the way in which T. recuvata handles photosynthesis. As is typical of the bromeliad family (Bromeliaceae), T. recuvata utilizes CAM photosynthesis. Instead of opening its stomata during the day, when high temperatures and baking sun would lead to unsustainable rates of water loss, T. recurvata opens its stomata at night, taking in CO2 while temperatures are more favorable. It then stores this CO2 as an organic acid that it can use later on the next day when the sun comes up. In doing so, T. recurvata can keep its stomata closed and save on water while still being able to synthesize the carbohydrates it needs.

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I think one of the main reasons T. recurvata doesn’t get the respect that many of its cousins receive is that it doesn’t put on a spectacular floral show when in bloom. Tiny purple to lavender petals just barely emerge from between bracts located a the tips of long flowers stalks. The flowers don’t last long and are quickly replaced by long, brown seed capsules. These capsules eventually burst open, releasing plenty of tiny seeds, each adorned with wispy filaments that help them take advantage of the slightest breeze. Though the seeds themselves are small and don’t show many adaptations for adhering to suitable substrates, I have found that those silky filaments tend to get matted up and stuck on whatever surface they land on. In this way, seeds at least have a chance to germinate on everything from twigs to power lines, and even other Tillandsias.

The reason this species earned the specific epithet ‘recurvata’ and the common name ‘ball moss’ has to do with both its growth habit and its propensity to grow on others of its own kind. Each leaf curls backward as it grows, giving individual plants a spherical shape. As more and more seedlings germinate on and around one another, these colonies can take on a massive, ball-like appearance. This has led many to classify this species as a parasite, however, this is not the case at all. It is wrongly assumed that these plants weaken the trees on which they grow and this is simply not the case.

Like many other epiphytes, T. recurvata likes a lot of sunlight. As such, plants tend to do better a the tops of trees or near the tips of branches. Certainly this can cause some degree of shading for the trees on which they grow, but this is insignificant considering how much a tree’s own branches and leaves shade those further down on the trunk. Also, T. recurvata are quick to move in on branches that have lost foliage or are already dead. This can often appear are is the plants have taken over the tree, causing it to die back. In reality, T. recurvata colonies are a merely a symptom of a tree already stressed by other factors. As the canopy starts to thin, more air plants are able to find suitable habitat for germination and growth. Trees covered in T. recurvata were already weak or dying, not the other way around.

In fact, evidence is showing that T. recurvata are actually an important source of nitrogen for the surrounding environment. Within their tissues, T. recurvata house specialized bacteria in the genus Pseudomonas, which are capable of fixing nitrogen directly from the atmosphere. In return for a place to live, these bacteria provide their air plant host with a nitrogen boost that would otherwise be unavailable. When T. recurvata detach from whatever they are growing on (something they frequently do in droves), they fall to the ground, rot, and enrich the soil with a shot of nitrogen. As such, these wonderful epiphytes are actually a boost to the growth of not only their hosts but many other plant species as well.

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Probably the most incredible feat of this species has been its conquest of the human environment. Throughout its range you can find T. recurvata thriving on man-made structures like power lines. For a species that gets all of its needs from the atmosphere, it is amazing how well T. recurvata is able to handle air pollution. Because it is so darn hardy, this air plant has caught the attention of more than one researcher. In fact, some are even looking at T. recurvata as a unique candidate for green roof construction in warmer climates.

All in all, this is one of the hardiest plants you are going to encounter in the Americas. One should look on at T. recurvata colonies with respect and admiration, not disgust and disdain. We fight species like this for all of the wrong reasons when in reality, we should be embracing them as both survivors and important components of ecosystem health. I hope this post has been able to do away with at least some of the misconceptions about this species. Three cheers for Tillandsia recurvata!

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

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

The Floating Bladderwort

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A carnivorous plant species that uses its radially arranged stolons like tiny pontoons to float at the waters surface may sound like something out of a science fiction novel. However, it is a very real strategy  adopted by one of the coolest carnivorous plants in North America. Utricularia inflata is one of the largest species of floating bladderwort on this continent and it is a species worth knowing.

Sometimes referred to as the swollen bladderwort, this species enjoys a native range that extends through much of the southeastern United States. For most of the year it exists in a state quite similar to other aquatic bladderworts. It has no true roots or leaves. Instead it produces a long, filiform stolon covered in tiny filaments that act as leaves with bladder traps situated at their tips. It sits in the water  column, gobbling up anything small and unfortunate enough to stumble into it.

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When flowering time approaches, these aquatic carnivores begin producing a different kind of stolon. Arranged like spokes on a wheel, the plant puts out swollen, air-filled stolons that float at the waters surface. These structures support the inflorescence. Flowers are bright yellow and resemble those of many other bladderwort species. Entire bodies of water can literally erupt in a sea of yellow bladderwort flowers when the right conditions present themselves.

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As mentioned, this species is carnivorous. It uses tiny bladder traps to suck in unsuspecting prey. Their diet is varied and includes pretty much anything that can fit into its bladder traps. One research paper reports both animal (rotifers, cladocerans, copepods, annelids, rhizopodeans, as well as small insects) and "plant" (Bacillariophyta, Chlorophyta, Cyanophyta, and Euglenophyta) prey.

Unfortunately these plants have been introduced far outside of their native range. In many areas they are becoming prevalent enough to be considered invasive. For instance, research done in the Adirondack Mountains of New York found that the presence of introduced populations of U. inflata caused significant changes in nutrient cycling, sediment chemistry, and overall net primary productivity.

This is a very neat species well worth a closer look. That being said, if you are a hobbyist such as myself, it is very important to remember that we should never release a species (no matter how cool it is) into areas where it isn't native.

Photo Credit: www.sarracenia.com, Dr. Mark Whitten, [3] [4]

Further Reading: [1] [2]

The Creeping Fuchsia

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Meet Fuchsia procumbens aka the creeping Fuchsia. This lovely plant is endemic to New Zealand where, sadly, it is threatened. In its native habitat, it is strictly a coastal species, prefering to grow in sandy soils. The  flowers are quite unlike most other members of the genus Fuchshia and they exhibit an interesting flowering strategy. 

Fuchsia procumbens produces 3 distinct flower forms, flowers with only  working male parts, flowers with only working female parts, and hermaphroditic flowers. One reason for this is to avoid self-pollination. The other reason may have something to do with energy costs. When growing conditions are less than stellar, the plant saves energy by producing male flowers. 

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Pollen is relatively cheap after all. When conditions improve, the plant may allocate more resources to female and hermaphroditic flowers. This strategy worries some botanists because it seems like some populations of F. procumbens only ever produce single sex flowers. After pollination, the flowers give way to bright red berries that are larger than the flowers themselves!

The most interesting thing about this species is, despite its apparent specificity in habitat preferences in the wild, it competes well with aggressive grasses, which has made it a very popular ground cover. As it turns out, its growing popularity in the garden trade may save this species from being placed on the endangered species list.

Photo Credits: [1] [2]

Further Reading: [1] [2]

The Smallest of the Giants

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There are a lot of cool ways to discover a new species but what about tripping over one? That is exactly how Rafflesia consueloae was found. Researchers from the University of the Philippines Los Baños were walking through the forest back in 2014 when one of them tripped over something. To their surprise, it was the bloom of a strange parasitic plant.

This was an exciting discovery because it meant that that strange family of holoparasitic plants called Rafflesiaceae just got a little bit bigger. Rafflesiaceae is famous the world over for the size of its flowers. Whereas the main body of plants in this family consists of tiny thread-like structures living within the tissues of forest vines, the flowers of many are huge. In fact, with a flower 3 feet (1 meter) in diameter, which can weigh as much as 24 lbs. (11 kg), Rafflesia arnoldii  produces the largest flower on the planet. This new species of Rafflesia, however, is a bit of a shrimp compared to its cousins.

In fact, R. consueloae produces the smallest flowers of the genus. Of the individuals that have been found, the largest flower clocked in at 3.83 inches (9.37 cm) in diameter. Needless to say, this was an exciting discovery and those responsible for it quickly set about observing the plant in detail. Cameras were set up to monitor flower development as well as to keep track of any animals that might pay it a visit. It turns out that, like its cousins, R. consueloae appears to be a specialist parasite on a group of vines in the genus Tetrastigma.

One of the unique characteristics of R. consueloae, other than its size, is the fact that its flowers don’t seem to produce any noticeable scent. This is a bit odd considering that its cousins are frequently referred to as “corpse flowers” thanks to the fact that they both look and smell like rotting meat. That is not to say that this species produces no scent at all. In fact, researchers noted that the fruits of R. consueloae smell a bit like coconut.

Its discoverers were quick to note that the discovery of such a strange parasitic plant in this particular stretch of forest is exciting because of the state of disrepair the forest is in. This region has suffered heavily from deforestation and fragmentation and it has long been thought that such specialized parasites like those in the genus Rafflesia could not persist after logging. As such, this discovery offers at least some hope that they may not be as sensitive as we once thought. Still, that does not mean that R. consueloae is by any means secure in its future.

To date, R. consueloae has only been found growing in two localities in Pantabangan, Phillippines. Though it is possible that more populations will be found growing elsewhere, its limited distribution nonetheless places it at high risk for extinction. Further habitat loss and the potential for anthropogenic forest fires are considerable threats to these plants and the hosts they simply can’t live without.

Despite plenty of observation, no one is quite sure how this species manages to reproduce successfully. Individual flowers are said to be either male or female but without a scent, its hard to say who or what pollinates them. Similarly, it still remains a mystery as to how R. consueloae (or any of its relatives for that matter) accomplish seed dispersal. Some small mammals were seen feeding on fruits but what happens after that is anyone’s guess. It seems like the various Rafflesiaceae still have many mysteries to be solved.

Photo Credit: [1]

Further Reading: [1]

 

The Peculiarly Tiny World of Buxbaumia Mosses

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Bug moss, bug-on-a-stick, humpbacked elves, elf-cap moss… Who knew there could be so many names for such tiny mosses. Despite their small stature, the mosses in the genus Buxbaumia have achieved something of a celebrity status to those aware of their existence. To find them, however, you need a keen eye, lots of patience, and a bit of luck.

Buxbaumia aphylla

Buxbaumia aphylla

Buxbaumia comprises something like 12 different species of moss scattered around much of the Northern Hemisphere as well as some parts of Australia and New Zealand. They are ephemeral in nature, preferring to grow in disturbed habitats where competition is minimal. More than one source has reported that they are masters of the disappearing act. Small colonies can arise for a season or two and then disappear for years until another disturbance hits the reset button and recreates the conditions they like.

Buxbaumia viridis

Buxbaumia viridis

I say you must have a keen eye and a lot of patience to find these mosses because, for much of their life, the exist on a nearly microscopic scale. Buxbaumia represents and incredible example of a reduction in body size for plants. Whereas the gametophytes of most mosses are relatively large, green, and leafy, Buxbaumia gametophytes barely exist at all. Instead, most of the “body” of these mosses consists of thread-like strands of cells called “protonema.” Though all mosses start out as protonema following spore germination, it appears that Buxbaumia prefer to remain in this juvenile stage until it comes time to reproduce.

Buxbaumia viridis

Buxbaumia viridis

Considering how small the protonemata are, there has been more than a little confusion as to how Buxbaumia manage to make a living. Early hypotheses suggested that these mosses were saprotrophs, living off of nutrients obtained from chemically digesting organic material in the soils. However, it is far more likely that these mosses rely heavily on partnerships with mycorrhizal fungi and cyanobacteria for their nutritional needs. It is thought that what little photosynthesis they perform is done via their protonema mats and developing sporophyte capsules.

Buxbaumia viridis

Buxbaumia viridis

Speaking of sporophytes, these are about the only way to find Buxbaumia in the wild. They are also the source of inspiration for all of those colorful common names. Compared to their gemetophyte stage, Buxbaumia sporophytes are giants. Fertilization occurs at some point in the fall and by late spring or early summer, the sporophytes are ready to release their spores. The size and shape of these capsules makes a lot more sense when you realize that they rely on raindrops for dispersal. When a drop impacts the flattened top of a Buxbaumia capsule, the spores are ejected into the environment and with any luck, will be carried off to another site suitable for growth.

Buxbaumia viridis

Buxbaumia viridis

I encourage you to keep an eye out for these plants. It goes without saying that data on population size and distribution is often lacking for such cryptic plants. Above all else, imagine how rewarding it would be to finally cross paths with this tiny wonders of the botanical world. Happy botanizing!

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


Maxipiñon: One of the Rarest Pines in the World

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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.

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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.

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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.

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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]

The Grafted Cactus Origin Story

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Many of you have undoubtedly met this interesting cactus before. Some  of you probably own one. Commonly referred to as 'Hibotan' or "moon  cactus," these are not a single species cactus but rather two different  cacti that have been grafted together.

The colorful top part is known scientifically as  Gymnocalycium mihanovichii. It is endemic to Paraguay and some provinces  of Argentina. In the wild it is not nearly this colorful. The specimens  sold in garden shops all over the world are actually mutant varieties that do not produce chlorophyll, thus revealing other pigments that are normally masked by green. The color of these mutants can range from  yellows to reds and even deep purples. Without chlorophyll, these mutants would normally die as seedlings.

The wild version of  Gymnocalycium mihanovichii  is a lot less coloreful.

The wild version of Gymnocalycium mihanovichii is a lot less coloreful.

Provided their host cactus is kept happy, mutant  Gymnocalycium mihanovichii  will flower.

Provided their host cactus is kept happy, mutant Gymnocalycium mihanovichii will flower.

At some point in time, someone got it in their head that they could graft these colorful mutants onto other species of cacti and perhaps they would survive. This is exactly what has happened. Interestingly enough, the bottom host cactus isn't even in the same genus as the moon cactus. Grafting is most often done on a species of Hylocereus (the same genus responsible for dragon fruit). How and why this host was chosen I do not know. Either way, armed with this knowledge, I hope you have gained a new found appreciation for these seemingly ubiquitous house plants.

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

Further Reading: [1]

Süßwassertang: A Fern Disguised as a Liverwort

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If you enjoy planted aquariums, you may have crossed paths with a peculiar little plant called Süßwassertang. It can be propagated by breaking off tiny pieces, which eventually grow into a tangled carpet of tiny green thalli. One could be excused for thinking that Süßwassertang was some sort of liverwort and indeed, for quite some time was marketed as such. That all changed in 2009 when it was revealed that this was not a liverwort at all but rather the gametophyte of a fern.

Despite its German name, Süßwassertang appears to have originated in tropical parts of Africa and Asia. It is surprisingly hard to find out any information about this plant outside of its use in the aquarium trade. The name Süßwassertang translates to “freshwater seaweed” and indeed, that is exactly what it looks like. The fact that this is actually the gametophyte of a fern may seem startling at first but when you consider what they must deal with in nature, the situation makes a bit more sense.

A  Süßwassertang gametophyte.  B  An antheridium, showing a cap cell ( cc ), ring cell ( rc ), and basal cell ( bc ).  Bar : 20 µm.  C  Developing lateral branches with rhizoids ( arrowhead ) and meristems ( m )  Bar : 0.2 mm.  D  Ribbon-like, branched gametophyte ( g ) of  L. spectabilis  bearing a young sporophyte ( sp )  Bar : 1 cm

A Süßwassertang gametophyte. B An antheridium, showing a cap cell (cc), ring cell (rc), and basal cell (bc). Bar: 20 µm. C Developing lateral branches with rhizoids (arrowhead) and meristems (m) Bar: 0.2 mm. D Ribbon-like, branched gametophyte (g) of L. spectabilis bearing a young sporophyte (sp) Bar: 1 cm

Fern gametophytes are surprisingly hardy considering their small size and delicate appearance. They are amazing in their ability to tolerate harsh conditions like drought and freezing temperatures. Because of this, fern gametophytes sometimes establish themselves in places that would be unfavorable for their sporophyte generation. For some, this means never completing their lifecycle. Others, however, seem to have overcome the issue by remaining in their gametophyte stage forever. Though no sexual reproduction occurs for these permanent gametophytes, they nonetheless persist and reproduce by breaking off tiny pieces, which grow into new colonies.

The sporophyte of a related species,  Lomariopsis marginata , demonstrating the usual epiphytic habit of this genus.

The sporophyte of a related species, Lomariopsis marginata, demonstrating the usual epiphytic habit of this genus.

This appears to be the case for Süßwassertang. Amazingly, despite a few attempts, no sporophytes have ever been coaxed from any gametophyte. It would appear that this is yet another species that has given up its sporophyte phase for an entirely vegetative habit. What is most remarkable is what the molecular work says about Süßwassertang taxonomically. It appears that this plant its nestled into a group of epiphytic ferns in the genus Lomariopsis. How this species evolved from vine-like ferns living in trees to an asexual colony of aquatic gametophytes is anyones’ guess but it is an incredible jump to say the least.

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

Further Reading: [1]

Gooey Pitcher Fluids

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There seems to be no end to the diversity of colors, shapes, and sizes exhibited by Nepenthes and their pitchers. These wonderful carnivorous plants grow these pitchers as a means of supplementing their nutritional needs as the habitats in which Nepenthes are found are lacking in vital nutrients like nitrogen. There are as many variations on the pitcher theme as there are Nepenthes but all function as traps in one form or another. How they trap insects is another topic entirely and some species have evolved incredible means of making sure prey does not escape. Some of my favorites belong to those species that employ sticky mucilage.

Arguably one of the most iconic of this type is Nepenthes inermis. This species is endemic to a small region of Sumatra and, to date, has only been found growing on a handful of mountain peaks in the western part of the country. The specific epithet ‘inermis’ is Latin for ‘unarmed’ as was given in reference to the bizarre upper pitchers of this plant. They look more like toilet bowls than anything carnivorous and indeed, they lack many of the features characteristic of other Nepenthes pitchers such as a peristome and a slippery, waxy coating on the inside of the pitcher walls.

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These may seem like minor details but consider the role these features play in other Nepenthes. A peristome is essentially a brightly colored, slippery lip that lines the outer rim of the pitcher mouth. Not only does this serve in attracting insect prey, it also aids in their capture. As mentioned, the peristome can be extremely slippery (especially when wet) so that any insect stumbling around on the rim is much more likely to fall in. Once inside, a waxy coating on the inside of some pitchers aids in keeping insects down. They simply cannot get purchase on the waxy walls and therefore cannot climb back out. So, for N. inermis to lack both features is a bit strange.

Another interesting feature of N. inermis pitchers is the highly reduced pitcher lid. It hasn’t disappeared completely but compared with other Nepenthes, this pitcher lid barely registers as one. For most Nepenthes, pitcher lids serve multiple functions. For starters, they keep the rain out. Nepenthes are msot at home in humid, tropical climates where rain is a daily force to be reckoned with. For many Nepenthes, rain not only dilutes the valuable digestive soup brewing within each pitcher, it can also cause them to overflow and dump their nutritious contents. Pitcher lids can also help in attracting prey. Like the peristome, they are often brightly colored but many also secrete nectar, which insects find irresistible. Lured in by the promise of food, some insects inevitably fall down into the pitcher below.

Looking into the pitcher of  Nepenthes inermis .

Looking into the pitcher of Nepenthes inermis.

Considering the importance of such structures, it becomes a little bit confusing why some Nepenthes have evolved away from this anatomy. The question then remains, why would a species like N. inermis no longer produce pitchers with these features? Amazingly, the answer actually lies within the pitcher fluid itself.

Tip over the upper pitchers of N. inermis and you will soon discover that they are filled with an extremely viscous mucilage. It is so viscous that some have reported that when the pitchers are held upside down, the mucilage within can form an unbroken stream of considerable length. Its the viscosity of this fluid that is the real reason that N. inermis is able to capture prey so easily. Insects lured to the traps can catch a drink of the nectar on the tiny lid. In doing so, some inevitably fall down into the pitcher itself.

The upper pitcher of the closely related  Nepenthes dubia .

The upper pitcher of the closely related Nepenthes dubia.

Instead of slippery walls or downward pointing hairs keeping the insects in, the viscous pitcher fluid quickly engulfs the struggling prey. Some have even suggested that the nectar secreted by the tiny lid has narcotic effects on visiting insects, however, I have not seen any data demonstrating this. Once caught in the fluid, insects easily slide their way down into the depths of the pitcher where they can be digested. This is probably why the pitchers are shaped like tiny toilet bowls; their shape allows for a large sticky surface area for insects to get stuck while prey that has already been captured is funneled down to where digestion and absorption takes place. In a way, these types of pitchers behave surprisngly similar to the sticky traps utilized by other carnivorous plants like sundews (Drosera spp.).

The viscous fluid also comes in handy during the frequent rains that blanket these mountains. As mentioned above, rain would quickly dilute most pitcher fluids but not when the pitcher fluid itself is more dense. Water sits on top of the viscous mucilage and when the pitchers become too heavy, they tip over. The water readily pours out but little if any of the pitcher fluid is lost in the process. It seems that species like N. inermis no longer fight the elements but rather have adapted to meet them head on. As such, they no longer have a need for a large pitcher lid.

Nepenthes jamban  takes the toilet bowl shape to the extreme.

Nepenthes jamban takes the toilet bowl shape to the extreme.

Nepenthes inermis is not alone in having evolved pitchers like this. Viscous pitcher mucilage is a trait shared by its closest relatives - N. dubia, N. flava, N. jacquelineae, N. jamban, N. talangensis, and N. tenuis, as well as even more distantly related species such as N. rafflesiana. Because prey capture is so important for the fitness of individuals, it is no wonder that so many different forms have evolved within this genus. In fact, many experts believe that variations in the way in which prey is captured and utilized is one of the main reasons why Nepenthes have undergone such a dramatic adaptive radiation.

Sadly, the uniqueness in form and function of these pitchers has landed many of these species on the endangered species list. As if habitat destruction wasn’t already pushing some to the brink, species like N. inermis are being poached at alarmingly unsustainable rates. Due to their limited distributions, most populations simply cannot recover from even moderate levels of harvesting. The silver lining in all of this is that many Nepenthes are extremely easy to grow and propagate provided their basic needs are met. As more and more folks enter into the carnivorous plant hobby, hopefully more and more people will be sharing seeds, cuttings, and tissue cultured materials. In doing so, we can hopefully reduce some of the pressures placed on wild populations.

Photos via Wikimedia Commons

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

A Herbaceous Conifer From the Triassic

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It is hard to make broad generalizations about groups of related organisms. There are always exceptions to any rule. Still, there are some “facts” we can throw around that seem to apply pretty well to specific branches on the tree of life. For instance, all of the gymnosperm lineages we share our planet with today are woody, relatively slow to reach sexual maturity, and are generally long-lived. This has not always been the case. Fossil discoveries from France suggest that in the past, gymnosperms were experimenting with a more herbaceous lifestyle.

The fossils in question were discovered in eastern France back in the 1800’s. The strata from which they were excavated dates back to the Middle Triassic, some 247 million years ago. Immortalized in these rocks were numerous spindly plants with strap-like leaves and a few branches, each ending in what look like tiny cones. Early interpretations suggested that these may represent an extinct lycopod, however, further investigation suggested something very surprising - a conifer with an herbaceous growth habit.

Indeed, thanks to even more scrutiny, it is now largely agreed upon that what was preserved in these rocks were essentially herbaceous conifers. The fossils were given the name Aethophyllum stipulare. They are wonderfully complete, depicting roots, shoots, leaves, and reproductive organs. Moreover, the way in which they were fossilized preserved lots of fine-scale anatomical details. Taken together, there are plenty of clues available that allow paleobotanists to say a lot about how this odd conifer made a living.

For starters, they were not very big plants. Not a single specimen has been found that exceeds 2 meters (6.5 ft) in height. The main stem of these conifers only seem to branch a couple of times. Cones were formed at the tips of the upper branches and not a single specimen has been found that depicts subsequent growth following cone formation. This suggests that Aethophyllum exhibited determinate growth, meaning that individuals grew to a certain size, reproduced, and did not continue to grow after that. Female cones were situated at the tips of the upper most branches and male cones were situated at the tips of lower shoots. The smallest reproductive individuals that have been unearthed are only 30 cm (11 in) in height, which suggests that Aethophyllum  was capable of reproducing within a few months of germination.

Artists reconstruction of  Aethophyllum stipulare

Artists reconstruction of Aethophyllum stipulare

Amazingly, researchers were also able to extract fossilized pollen and seeds from some of the Aethophyllum cones. The pollen itself is saccate, much like what we see in many extant conifers. By comparing the morphology of the pollen extracted from the cones to other fossil pollen records, researchers now feel confident that Aethophyllum is the source of pollen grains discovered in sediments from western, central, and southern Europe, Russia, Northern Africa, and China, suggesting that Aethophyllum was pretty wide spread during the Middle Triassic. Aethophyllum seeds were small, ellipsoid, and were not winged, likely germinating a short distance from the parent.

The stems of Aethophyllum are interesting in the own right. Thanks to their preservation, cross sections have been made and they reveal that these plants only ever produced secondary tracheids and primary xylem. The only place on the plant where any signs of woody secondary xylem occur are at the base of the cones. This adds further confirmation that Aethophyllum was herbaceous at the onset of sexual maturity.

Another intriguing aspect of the stem is the presence of numerous large air spaces within the stem pith. Today, this anatomical feature is present in plants like bamboo, Equisetum, and the flowering stalks of Agave, all of which exhibit alarmingly fast growth rates for plants. This suggests that not only did Aethophyllum reproduce early in its life, it also likely grew extremely fast.

1. Smallest fertile plant in the Grauvogel and Gall collections, with two stems extending from the root, and terminal ovulate cone (OC) on one branch (scale bar=10 cm). 2. Cross-section of stem in the Grauvogel and Gall collections showing cauline bundles with scanty wood (at left, top and right) surrounding large pith with large, aerenchymatous lacunae and interspersed pith parenchyma cells. Vascular cambium, phloem, and more peripheral tissues are not preserved (scale bar=200 μm). 3.Seedling in the Grauvogel and Gall collections showing primary root (R), cotyledons (C) and stem (S) with apically borne leaves (scale bar=10 cm).  Quoted from SOURCE

1. Smallest fertile plant in the Grauvogel and Gall collections, with two stems extending from the root, and terminal ovulate cone (OC) on one branch (scale bar=10 cm). 2. Cross-section of stem in the Grauvogel and Gall collections showing cauline bundles with scanty wood (at left, top and right) surrounding large pith with large, aerenchymatous lacunae and interspersed pith parenchyma cells. Vascular cambium, phloem, and more peripheral tissues are not preserved (scale bar=200 μm). 3.Seedling in the Grauvogel and Gall collections showing primary root (R), cotyledons (C) and stem (S) with apically borne leaves (scale bar=10 cm). Quoted from SOURCE

Mature Aethophyllum aren’t the only fossils available either. Many seedlings have been discovered in close proximity to the adults. Seedlings were also exquisitely preserved, depicting hypocotyl, a primary root system, two two-veined cotyledons, and a short stem with four-veined leaves arranged in a helix. The fact that seedlings and adults were found in such close proximity lends to the idea that Aethophyllum populations were made up of multi-aged stands, not unlike some of the early successional plants we find in disturbed habitats today.

The sediments in which these plants were fossilized can also tell us something about the habitats in which Aethophyllum grew. The rock layers are made up of a mix of sediments typical of what one would find in a flood plain or delta. Also, Aethophyllum aren’t the only plant remains discovered. Many species known to grow in regularly disturbed, flood-prone habitats have also been found. Taken together these lines of evidence suggest that Aethophyllum was similar to what we would expect from herbaceous plants growing in similar habitats today. They grew fast, reproduced early, and had to jam as many generations in before the next flood ripped through and hit the reset button.

Aethophyllums small size, lack of wood, and rapid growth rate all point to a ruderal lifestyle. Today, this niche is largely filled by angiosperms. No conifers alive today can claim such territories. The discovery of Aethophyllum demonstrates that this was not always the case. The fact that pollen has been found far outside of France suggests that this ruderal lifestyle worked quite well for Aethophyllum.

The terrestrial habitats of the Middle Triassic were dominated by the distant relatives of modern day ferns, lycophytes, and gymnosperms. Needless to say, it was a very different world than anything that we are familiar with today. However, that does not mean that the pressures of natural selection were necessarily different. Aethophyllum is evidence that specific selection pressures, in this case regular flood disturbance, select for similar traits in plants through time. Why Aethophyllum went extinct is anyone’s guess. Despite how well they have been preserved, there is still a lot of mystery surrounding this plant.

Photo Credit: [1]

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



The Celery-Topped Conifers

I am only just starting to fully appreciate the diversity in form and habit exhibited by the gymnosperm lineages alive today. What I once thought of as a unidimensional group of plants is proving to be wonderfully diverse, despite being overshadowed by the angiosperms. For instance, imagine my surprise when I first laid eyes on a member of the genus Phyllocladus.

At first glance, these weird conifers look more like a broad-leaf angiosperm. This similarity is superficial, of course. Before we get to why they look the way they do, it is worth considering this group from a as a whole. The genus Phyllocladus comprises roughly 5 species spread out among New Zealand, Tasmania, and Malesia. They are somewhat variable in form but usually settle out somewhere between a good sized shrub and a medium sized tree. Where exactly this genus of oddball gymnosperms fits on the tree of life is subject to some debate.

Phyllocladus aspleniifolius

Phyllocladus aspleniifolius

Phyllocladus trichomanoides

Phyllocladus trichomanoides

For many years after its initial description, Phyllocladus was placed in a family of its own - Phyllocladaceae. Subsequent molecular work has only managed to add to the confusion. Despite its unique morphological characteristics, some authors feel this genus fits nicely into the family Podocarpaceae. At least one other study suggests that it doesn’t belong in Podocarpaceae but rather is situated as sister to the family. By the looks of it, this will not be cleared up any time soon. So, for now, let’s focus in on why these plants are so strange.

For starters we have the “leaves.” I place the word ‘leaves’ in quotes because they are not true leaves. The correct term for these structures are phylloclades (hence the generic name). A phylloclade is a flattened projection of a branch that takes on the form and function of a leaf. What we know of as leaves have been greatly reduced in the genus Phyllocladus. If you want to see them, you must look closely at the tips of the phylloclades. Early on in their development, the leaves exist as tiny brown scales. These scales are gradually lost over time as they serve no function for the plant.

Phyllocladus alpinus

Phyllocladus alpinus

Phyllocladus hypophyllus

Phyllocladus hypophyllus

Though no one has tested this directly (that I am aware of), the evolution of phylloclades over leaves likely has to do with energy conservation in one form or another. Why produce stems and leaves when you can co-opt stem-like structures to do the work for you? Oddly enough, some suggest that to consider them stems in the truest sense of the word is erroneous. Morphologically speaking, they share traits that are intermediate between branches and stems. However, I am going to need to do more homework before I feel comfortable elaborating on this point.

Only when it comes time for reproduction does their place among the gymnosperms become readily apparent, that is before the ovules are fertilized. All members of the genus Phyllocladus produce cones. Male cones are tiny, cylindrical structures located at the ends of their side branches whereas female cones are clustered into groups along the axils or margins of the phylloclades. Once fertilized, however, these plants offer another point of confusion for the casual observer.

The fleshy “fruits” of  Phyllocladus aspleniifolius

The fleshy “fruits” of Phyllocladus aspleniifolius

Phyllocladus is yet another genus of conifers that has converged on a fruit-like seed dispersal strategy. As the seed cones mature, the scales gradually swell and become berry-like. Poking out of the bright red and/or white aril is a single seed. These fleshy arils function in a similar way to fruit in that they attract birds, which then consume them, dispersing the seeds later on in their feces.

Another intriguing aspect of their morphology occurs below ground. The roots of this genus form nodules, which provide a home for bacteria that specializing in fixing atmospheric nitrogen. In return for a home and some carbohydrates from photosynthesis, these bacteria pay these trees with nitrogen that would otherwise be unavailable. Pretty remarkable stuff for a such an esoteric group of conifers!

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Photo Credits: [1] [2]

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

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.

Nuytsia_floribunda_-_The_Australian_Mistletoe.jpg

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

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