Goblin's Gold: the story of a luminous moss

Photo by Alpsdake licensed under CC BY-SA 4.0

Luminous moss, dragon’s gold, goblin’s gold… when a moss has this many common names, you know it must catch the eye. Indeed, Schistostega pennata might just be one of the most dazzling of mosses around, that is provided you know where and how to look for it.

Let’s begin with a brief introduction. Goblin’s gold is the only member of both its genus (Schistostega) and family (Schistostegaceae). Despite its unique taxonomic position, it is nonetheless a widespread species, growing naturally throughout many temperate regions of the Northern Hemisphere.

When fully grown, the gametophyte stage of goblin’s gold sort-of resembles a tiny, green, semi-translucent feather. Small spore capsules are borne on the spindly stalk of the sporophyte and the resulting spores are said to be quite sticky. Instead of relying on wind to disperse its propagules, golbin’s gold utilizes animals. The spores are sticky enough that they get glom onto any insects or other small animals that brush up against them.

The mature gametophyte of Schistostega pennata. Photo by HermannSchachner licensed under Public Domain

None of this, however, gives a hint as to how it earned all of those colorful names. To find that out, one must be ready to brave dark, damp spaces like caves. You see, though it can grow in more open habitats, you are most likely to encounter goblin’s gold in dark crevices or under overhangs. It has been said that goblin’s gold does not compete well with other plants in most habitats, but that doesn’t mean it doesn’t have a few tricks up its stems that give it an edge in other types of habitats.

For most plants, caves and other dark places are a no go. They simply can’t get enough light to survive. Such is not the case for goblin’s gold. Instead of trying to compete with more aggressive vegetation, goblin’s gold occupies deeply shaded habitats that few other plants can. It owes its shade-tolerant abilities to a stage of its development most of us rarely think about, let alone notice.

Photo by Jymm licensed under CC BY-SA 4.0

When a moss spore germinates, it doesn’t immediately look like what we would recognize as a moss. Instead, it grows into thread-like, multicellular fillaments called a “protonema.” You can think of this as the juvenile stage of the gametophyte. The protonema spreads outward as it grows, gradually producing hormones and other growth regulators that will control the development of the mature gametophyte. Because goblin’s gold grows in such dark habitats, it can’t afford to grow its gametophyte anywhere. To grow long enough to reproduce, it has to find spots where there is enough light to complete its lifecycle.

This is where the protonema comes in. In much the same why that fungal hyphae fan out into the soil in search of food to decompose, goblin’s gold protonema fan out over the damp substrate, searching for spots where enough light filters through to fuel growth. Luckily, the protonema can make do with much less light that the mature gametophyte, which also happens to be how this tiny moss earned so many interesting nicknames.

When grown in deep shade, the protonema of goblin’s gold develops a layer of lens-shaped cells on its surface. The opposite side of each cell narrows to a cone. When light, no matter how weak, strikes these lens cells, the curvature focuses the light down into the cell so that it is concentrated into the tip at the bottom. Being able to sense the direction of the light, the chloroplasts within each cell can actually move around so that they are always in a position that maximizes their exposure. Through this process, each cell is able to concentrate what little light is available so that they can photosynthesize in light so low that nearly all other plants will starve.

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The light concentrating mechanism of the goblin’s gold protonema happens to have a wonderful and stunning side effect. As light enters the lens, small amounts of it are refracted around the cell. When that refracted light mixes with the green light that isn’t absorbed by the chloroplasts, it bounces back into the environment, giving the whole protonemal mat a green florescent glow when viewed in just the right way.

By being able to make use of what little light finds its way into these dark habitats, goblin’s gold can grow largely free of competition. Also, the protonema itself is capable of asexual reproduction so colonies can grow to epic proportions in dark areas, only producing mature gametophytes in a few spots. Interestingly, there appears to be some plasticity to this light-concentrating habit as well. When observing goblin’s gold protonema that develop under high light conditions, researchers have found that they do not develop lens shaped cells and therefore are not capable of reflecting light in the same way.

Humans have known about this moss for centuries, even if they didn’t understand the mechanisms that cause it, and that is why this wonderfully unique species has earned so many common names.

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

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

The Peculiarly Tiny World of Buxbaumia Mosses

Photo by Tab Tannery licensed under CC BY-NC-SA 2.0

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. Photo by Bernd Haynold licensed under CC BY-SA 4.0

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. Photo by BerndH licensed under CC BY-SA 3.0

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. Photo by Bernd Haynold licensed under CC BY-SA 4.0

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. Photo by Bernd Haynold licensed under CC BY-SA 3.0

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. Photo by BerndH licensed under CC BY-SA 3.0

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]

An Introduction to Hornworts

Anthoceros sp. Photo by Bramadi Arya licensed under CC BY-SA 4.0

When was the last time you thought about hornworts? Have you ever thought about hornworts? If you answered no, you aren’t alone. Despite their global distribution, these tiny plants receive hardly any attention and that is a shame. Hornworts (Anthocerotophyta) have been around for a very long time. In fact, it is likely that they were some of the first plants to colonize the land roughly 300 - 400 million years ago.

To be fair, hornworts aren’t known for their size. They are generally small plants, though their colonies can form impressive mats. To find them, one must try looking in and among rocks, bare patches of soil, or pretty much anywhere enough moisture builds up to supply their needs. They tend to enjoy nutrient-poor substrates but I would hesitate to say that with any certainty. No matter where you live, from the tundra to the tropics, there is probably a hornwort native to your neck of the woods.

Dendroceros sp. Photo by J.Ziffer licensed under public domain

How many different species of hornwort there are is apparently the subject of some debate. Some authors recognize upwards of 300 species whereas others suggest the real number hangs somewhere around 150. Regardless of the exact numbers, hornworts belong to one of six genera: Anthoceros, Dendroceros, Folioceros, Megaceros, Notothylas and Phaeoceros. Fun fact, the suffix ‘ceros’ at the end of each genus is derived from the Latin word for ‘horn.’

The reason they are called hornworts is because of their reproductive structures or “sporophytes.” Similar to their moss and liverwort cousins, hornworts undergo an alternation of generations in order to reproduce sexually. The green gametophytes house the sexual organs - antheridia if they are male and archegonia if they are female. After fertilization, a sporophyte begins to grow, which will go on to produce and disseminate spores. However, the way in which the hornwort sporophyte forms is a bit different from what we see in mosses and liverworts.

Alternation of generations in hornworts. Photo by Mariana Ruiz (LadyofHats) licensed under public domain

Upon fertilization, the zygote begins to divide into a bulbous mass of cells affectionately referred to as "the foot.” This foot remains within the gametophyte throughout the lifetime of the hornwort, depending on the gametophyte for water and nutrients. Even more peculiar is the the fact that the growing point of the sporophyte is at the base rather than the tip. As such, the horn of each hornwort could continue to grow upwards until it is damaged in some way.

The horn itself is an amazing structure. Whereas the outside layers of tissue are merely structural, the internal tissues differentiate into two different types - spores and pseudo-elaters. Pseudo-elaters expand and contract as humidity fluctuates so as the sporophyte splits to release the spores, the pseudo-elaters dehydrate and snap like tiny spore catapults, thus aiding in their dispersal.

Megaceros flagellaris. Photo by Dr. Scott Zona licensed under CC BY-NC 2.0

Of course, reproduction is the main goal but to get to that point, hornworts must grow and mature. How they manage to survive is incredible because it is a reminder that what are often thought of as “primitive” plants are actually far more advanced than we give them credit for. The main body of the hornwort gametophyte is a thin layer of cells that spread out to form a tiny, green mat. This is the structure you are most likely to encounter.

Inside each cell is a single chloroplast. In most hornworts, the chloroplast does not exist in isolation. Instead, it is fused with other organelles into a structure called a “pyrenoid.” The pyrenoid functions as both a center for photosynthesis and a food storage organ. This is unique as it relates to terrestrial plants but quite common in algae. Another odd fact about hornwort anatomy are the presence of tiny cavities scattered throughout their tissues. These cavities form as clusters of hornwort cells die. They then fill with a special mucilage that appears to invite colonization by nitrogen-fixing cyanobacteria. The cyanobacteria set up shop within the cavities and provides the hornwort with supplemental nitrogen in return for a place to live.

Anthoceros agrestis photo by BerndH licensed under CC BY-SA 3.0

Cyanobacteria aren’t the only organisms to have partnered with hornworts either. Mycorrhizal fungi also enter into the picture. A study done back in 2013 actually found that a wide variety of fungi will partner with hornworts which suggests that this symbiotic relationship is much more ancient and versatile than we once thought. Fungi cluster around parts of the gametophyte that produce root-like structures called “rhizoids,” offering nutrients in return for carbohydrates.

All in all, I think it is safe to say that hornworts are remarkable little plants. Though they can sometimes be difficult to find and properly identify, they nonetheless offer plenty of inspiration for the botanically inclined mind. We can all do better by tiny plants like the hornworts. They have been on land for an incredible amount of time and they definitely deserve our respect and admiration.

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

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

Tropical Ferns in Temperate North America

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All plants undergo some form of alternation of generations. It is the process in which, through reproduction, they cycle between a haploid gametophyte stage and a diploid sporophyte stage. In ferns and lycophytes, this alternation of generations has been taken to the extreme. Instead of the sporophyte relying on the gametophyte for sustenance, the two generations are physically independent and thus separated from one another. In a handful of fern genera here in North America, this has led to some intriguing and, dare I say, downright puzzling distributions.

The presence of a small handful of tropical fern genera in temperate North America has generated multiple scientific investigations since the early 1900's. However, as is constantly happening in science, as soon as we answer one question, seemingly infinite more questions arise. At the very least, the presence of these ferns in temperate regions offers us a tantalizing window into North America’s ancient past.

To say any of these ferns offer the casual observer much to look at would be a bit of an exaggeration. They do not play out their lives in typical fern fashion. These out-of-place tropical ferns exists entirely as asexual colonies of gametophytes, reproducing solely by tiny bundles of cells called gemmae. What's more, you will only find them tucked away in the damp, sheltered nooks and crannies of rocky overhangs and waterfalls. Buffered by unique microclimates, it is very likely that these fern species have existed in these far away corners for a very, very long time. The last time their respective habitats approached anything resembling a tropical climate was over 60 million years ago. Some have suggested that they have been able to hang on in their reduced form for unthinkable lengths of time in these sheltered habitats. Warm, wet air gets funneled into the crevices and canyons where they grow, protecting them from the deep freezes so common in these temperate regions. Others have suggested that their spores blew in from other regions around the world and, through chance, a few landed in the right spots for the persistence of their gametophyte stages.

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The type of habitat you can expect to find these gametophytes.

Aside from their mysterious origins, there is also the matter of why we never find a mature sporophyte of any of these ferns. At least 4 species in North America are known to exist this way - Grammitis nimbata, Hymenophyllum tunbridgense, Vittaria appalachiana, and a member of the genus Trichomanes, most of which are restricted to a small region of southern Appalachia. In the early 1980's, an attempt at coaxing sporophyte production from V. appalachiana was made. Researchers at the University of Tennessee brought a few batches of gametophytes into cultivation. In the confines of the lab, under strictly controlled conditions, they were able to convince some of the gametophytes to produce sporophytes. As these tiny sporophytes developed, they were afforded a brief look at what this fern was all about. It confirmed earlier suspicions that it was indeed a member of the genus Vittaria, or as they are commonly known, the shoestring ferns. The closest living relative of this genus can be found growing in Florida, which hints at a more localized source for these odd gametophytes. However, both physiology and subsequent genetic analyses have revealed the Appalachian Vittaria to be a distinct species of its own. Thus, the mystery of its origin remains elusive.

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In order to see them for yourself, you have to be willing to cram yourself into some interesting situations. They really put the emphasis on the "micro" part of the microclimate phenomenon. Also, you really have to know what you are looking for. Finding gametophytes is rarely an easy task and when you consider the myriad other bryophytes and ferns they share their sheltered habitats with, picking them out of a lineup gets all the more tricky. Your best bet is to find someone that knows exactly where they are. Once you see them for the first time, locating other populations gets a bit easier. The casual observer may not understand the resulting excitement but once you know what you are looking at, it is kind of hard not to get some goosebumps. These gametophyte colonies are a truly bizarre and wonderful component of North American flora.

Photo Credit: [1] [2]

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

Meet The Powder Gun Moss

I get very excited when I am able to identify a new moss. This is mainly due to the fact that moss ID is one of my weakest points. I was sitting down on a rock the other day taking a break from vegetation surveys when I looked to my right and saw something peculiar. The area was pretty sloped and there was some exposed soil in the vicinity. Covering some of that soil was what looked like green fuzz. Embedded in that fuzz were these strange green urns.

I busted out my hand lens and got a closer look. This was definitely a moss but one I had never seen before. The urns turned out to be capsules. Later, a bit of searching revealed this to be a species of moss in the genus Diphyscium. This genus is the largest within the family Diphysciaceae and here in North America, we have two representatives - D. foliosum and D. mucronifolium.

These peculiar mosses have earned themselves the common name 'powder gun moss.' The reason for this lies in those strange sessile capsules. Unlike other mosses that send their capsules up on long, hair-like seta in order to disperse their spores on the faintest of breezes, the Diphyscium capsules remain close to the ground. In lieu of wind, a powder gun moss uses rain. In much the same way puffball mushrooms harness the pounding of raindrops, so too do the capsules of the powder gun moss. Each raindrop that hits a capsule releases a cloud of spores that are ejected into an already humid environment full of germination potential.

Luckily for moss lovers like myself, the two species of Diphyscium here in North America tend to enjoy very different habitats. This makes a positive ID much more likely. D. foliosum prefers to grow on bare soils whereas D. mucronifolium prefers humid rock surfaces. Because of this distinction, I am quite certain the species I encountered is D. foliosum. And what a pleasant encounter it was. Like I said, it isn't often I accurately ID a moss so this genus now holds a special place in my mind.

Further Reading: [1] [2]

Growing Ferns

I am finally having some success intentionally growing ferns from spores. I collected and sowed spores from some interrupted ferns (Osmunda claytoniana) over the summer. They have been hanging out as gametophytes for months now and some are finally starting to grow sporophytes. Here is how it worked for me:

I kept my eye on a batch of adult plants this summer. Once their fertile fronds developed I would flick them every now and then to see if they were releasing spores. Once I saw that they were I shook the fronds over some paper to collect the spores. I then took some old potting soil and sterilized it with boiling distilled water. I use old takeout containers because they are small and have clear lids that form a seal which keeps the humidity high.

Once the soil was cool I sprinkled the spores over it and then placed it on a shelf where it gets a small amount of ambient light every day. The rest they did themselves. You just have to remember to check on them and keep the humidity quite high because they can dry out really fast. They seemed stuck as gametophytes for months. I just noticed the start of these sporophytes the other day.