What Is a Plant, and Why Should I Care? part three

“If it wasn’t for the plants, and if it wasn’t for the invertebrates, our ancestors’ invasion of land could never have happened. There would have been no food on land. There would have been no ecosystems for them to populate. So really the whole ecosystem that Tiktaalik and its cousins were moving into back in the Devonian was a new ecosystem. … This didn’t exist a hundred million years before – shallow fresh water streams with soils that are stabilized by roots. Why? Because it took plants to do that – to make the [habitats] in the first place. So really plants, and the invertebrates that followed them, made the habitats that allowed our distant relatives to make the transition from life on water to life on land.” – Neil Shubin, author of Your Inner Fish, in an interview with Cara Santa Maria on episode 107 of her podcast, Talk Nerdy To Me

Plants were not the first living beings to colonize land – microorganisms have been terrestrial for what could be as long as 3.5 billion years, and lichens first formed on rocks somewhere between 550 and 635 million years ago – however, following in the footsteps of these other organisms, land plants paved the way for all other forms of terrestrial life as they migrated out of the waters and onto dry land.

The botanical invasion of land was a few billion years in the making and is worth a post of its own. What’s important to note at this point, is that the world was a much different place back then. For one, there was very little free oxygen. Today’s atmosphere is 21% oxygen; the first land plants emerged around 470 million years ago to an atmosphere that was composed of a mere 4% oxygen. Comparatively, the atmosphere back then was very carbon rich. Early plants radiated into numerous forms and spread across the land and, through processes like photosynthesis and carbon sequestration, helped to dramatically increase oxygen levels. A recent study found that early bryophytes played a major role in this process. The authors of this study state, “the progressive oxygenation of the Earth’s atmosphere was pivotal to the evolution of life.”

A recreation of a Cooksonia species - one of many early land plants. (photo credit: wikimedia commons)

A recreation of a Cooksonia species – one of many early land plants (photo credit: wikimedia commons)

The first land plants looked very different compared to the plants we are used to seeing today. Over the next few hundred million years plants developed new features as they adapted to life on land and to ever-changing conditions. Roots provided stability and access to water and nutrients. Vascular tissues helped transport water and nutrients to various plant parts. Woody stems helped plants reach new heights. Seeds offered an alternative means of preserving and disseminating progeny. Flowers – by partnering with animal life – provided a means of producing seeds without having to rely on wind, water, or gravity. And that’s just scratching the surface. Rooted in place and barely moving, if at all, plants appear inanimate and inactive, but it turns out they have a lot going on.

But what is a plant again? In part one and two, we listed three major features all plants have in common – multicellularity, cell walls composed of cellulose, and the ability to photosynthesize – and we discussed how being an autotroph (self-feeder/producer) sets plants apart from heterotrophs (consumers). Joseph Armstrong writes in his book, How the Earth Turned Green, “photosynthetic producers occupy the bottom rung of communities.” In other words, “all modern ecosystems rely upon autotrophic producers to capture energy and form the first step of a food chain because heterotrophs require pre-made organic molecules for energy and raw materials.”

So, why should we care about plants? Because if it wasn’t for them, there wouldn’t be much life on this planet to speak of, including ourselves.

Plants don’t just provide food though. They provide habitat as well. Plus they play major roles in the cycling of many different “nutrients,” including nitrogen, phosphorous, carbon, sulfur, etc. They are also a major feature in the water cycle. It is nearly impossible to list the countless, specific ways in which plants help support life on this planet, and so I offer two examples: moss and dead trees.

The diminutive stature of mosses may give one the impression that they are inconsequential and of little use. Not so. In her book, Gathering Moss, Robin Wall Kimmerer describes how mosses support diverse life forms:

There is a positive feedback loop created between mosses and humidity. The more mosses there are, the greater the humidity. More humidity leads inexorably to more mosses. The continual exhalation of mosses gives the temperate rain forest much of its essential character, from bird song to banana slugs. … Without mosses, there would be fewer insects and stepwise up the food chain, a deficit of thrushes.

Mosses are home to numerous invertebrate species. For many insects, mosses are a place to deposit their eggs and, consequentially, a place for their larvae to mature into adults. Banana slugs traverse the moss feeding on “the many inhabitants of a moss turf, and on the moss itself.” In the process they help to disperse the moss.

Moss is used as a nesting material by various species of birds, as well as squirrels, chipmunks, voles, bears, and other animals. Patches of moss can also function as “nurseries for infant trees.” In some instances, mosses inhibit seed germination, but they can also help protect seeds from drying out or being eaten. Kimmerer writes, “a seed falling on a bed of moss finds itself safely nestled among leafy shoots which can hold water longer than the bare soil and give it a head start on life.”

moss as nurse plant

Virtually all plants, from the tiniest tufts of grass to the tallest, towering trees have similar stories to tell about their interactions with other living things. Some have many more interactions than others, but all are “used” in some way. And even after they die, plants continue to interact with other organisms, as is the case with standing dead trees (a.k.a. snags).

In his book, Welcome to Subirdia, John Marzluff explains that when “hole creators” use dead and dying trees, they benefit a host of “hole users:”

Woodpeckers are natural engineers whose abandoned nest and roost cavities facilitate a great diversity of life, including birds, mammals, invertebrates, and many fungi, moss, and lichens. Without woodpeckers, birds such as chickadees and tits, swallows and martins, bluebirds, some flycatchers, nuthatches, wood ducks, hooded mergansers, and small owls would be homeless.

As plants die, they continue to provide food and habitat to a variety of other organisms. Eventually they are broken down to their most rudimentary components, and their nutrients are taken up and used by “new life.” Marzluff elaborates on this process:

Much of the ecological web exists out of sight – underground and in rotting wood. There, molds, bacteria, fungi, and a world of invertebrates convert the last molecules of sun-derived plant sugar to new life. These organisms are technically ‘decomposers,’ but functionally they are among the greatest of creators. Their bodies and chemical waste products provide us with an essential ecological service: soil, the foundation of terrestrial life.

Around 470 million years ago, plants found their way to land. Since then life of all kinds have made land their home. Plants helped lead the way. Today, plants continue their long tradition of supporting the living, both in life and in death.

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Dung Moss (Revisited)

This is a revised version of a post that was originally published on January 14th, 2015. It includes excerpts from a chapter entitled, “Portrait of Splachnum,” in the book, Gathering Moss, by Robin Wall Kimmerer.

Certain plants, like corpse flowers and carrion flowers, emit foul odors when they bloom. The scent is akin to the smell of rotting flesh, hence their common names. The purpose of this repugnant act is to attract a specific group of pollinators: flies, carrion beetles, and other insects that are attracted to gross things. Though this particular strategy is rare, these aren’t the only plants that employ stinky smells to recruit such insects to aid in reproduction and dissemination. Consider dung mosses.

No moss is more fastidious in its choice of habitats than Splachnum. Absent from the usual mossy haunts, Splachnum is found only in bogs. Not among the commoners like Sphagnum that build the peaty hummocks, not along the margins of the blackwater pools. Splachnum ampullaceum occurs in one, and only one, place in the bog. On deer droppings. On white-tailed deer droppings. On white-tailed deer droppings which have lain on the peat for four weeks. In July.

At least three genera (SplachnumTetraplodon, and Tayloria) in the family Splachnaceae include species that go by the common name, dung moss. All Splachnum and Tetraplodon species and many species in the genus Tayloria are entomophilous. Entomophily is a pollination strategy in which pollen or spores are distributed by insects. Compare this to anemophily, or wind pollination, which is the common way that moss spores are distributed. In fact, dung mosses are the only mosses known to exhibit entomophily.

Dung Moss (photo credit: wikimedia commons)

Dung Moss (photo credit: wikimedia commons)

Before we go too much further, it’s important to understand how mosses differ from other plants. Mosses are in a group of non-vascular and non-flowering plants called bryophytes. Vascular tissues are the means by which water and nutrients are transported to and from plant parts. Lacking vascular tissues, water and nutrients are simply absorbed through the leaves and stems of mosses, which is why mosses are typically petite and prefer moist environments. Mosses also lack true roots and instead have rhizoids – threadlike structures that anchor the plants to their substrate of choice (such as dung).

Another major distinction between bryophytes and other plants is that bryophytes spend most of their life cycle as a haploid gametophyte rather than a diploid sporophyte. In most plants, the haploid gametophytes are the sperm (pollen) and egg cells; the sporophyte is everything else. In mosses, the familiar green, leafy structure is actually the gametophyte. The gametophyte houses sperm and egg cells, and when the egg is fertilized by sperm it forms a zygote that develops into the sporophyte structure which extends above the leafy gametophyte. A capsule at the top of the sporophyte contains spores which are eventually released and, upon finding themselves on a suitable substrate in a hospitable environment, germinate to produce new plants. The spore then is comparable to a seed in vascular, seed-bearing plants.

photo credit: wikimedia commons

photo credit: wikimedia commons

As stated earlier, the spores of most mosses are distributed by wind. Dung mosses, on the other hand, employ flies in the distribution of their spores. They attract the flies by emitting scents that only flies can love from an area on the capsule of the sporophyte called the apophysis. This area is often enlarged and brightly colored in yellow, magenta, or red, giving it a flower-like appearance which acts as a visual attractant. The smells emitted vary depending on the type of substrate a particular species of dung moss inhabits. Some dung mosses grow on the dung of herbivores and others on the dung of carnivores. Some even prefer the dung of a particular group of animals; for example, a population of Tetraplodon fuegiensis was found to be restricted to the feces and remains of foxes. However, dung is not the only material that dung mosses call home. Certain species grow on rotting flesh, skeletal remains, or antlers.

Splachnum ampullaceum inhabits the droppings of white-tailed deer. Had a wolf or coyote followed the scent of the deer into the bog, its droppings would been colonized by S. luteum. The chemistry of carnivore dung is sufficiently distinct from that of herbivores to support a different species. … Moose droppings have their own loyal follower. The family to which Splachnum belongs includes several other mosses with an affinity for animal nitrogen. Tetraplodon and Tayloria can be found on humus, but primarily inhabit animal remains such as bones and owl pellets. I once found an elk skull lying beneath a stand of pines, with the jawbone tufted with Tetraplodon.

Yellow Moosedung Moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit: www.eol.org)

Yellow moosedung moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit: www.eol.org)

The set of circumstances that converge to bring Splachnum into the world is highly improbable. Ripening cranberries draw the doe to the bog. She stands and grazes with ears alert, flirting with the risk of coyotes. Minutes after she has paused, the droppings continue to steam. … The droppings send out an invitation written in wafting molecules of ammonia and butyric acid. Beetles and bees are oblivious to this signal, and go on about their work. But all over the bog, flies give up their meandering flights and antennae quiver in recognition. Flies cluster on the fresh droppings and lap up the salty fluids that are beginning to crystallize on the surface of the pellets. Gravid females probe the dung and insert glistening white eggs down into the warmth. Their bristles leave behind traces from their earlier foraging trips among the day’s dung, delivering spores of Splachnum on their footprints.

The spores of dung mosses are small and sticky. When a fly visits these plants, the spores adhere to its body in clumps. The fly then moves on to its substrate of choice to lay its eggs, and the spores are deposited where they can germinate and grow into new moss plants. Flies that visit dung mosses receive nothing in return for doing so, but instead are simply “tricked” into disseminating the propagules. The story is similar with corpse flowers and carrion flowers; flies are drawn in by the smells and recruited to transmit pollen while receiving no nectar reward for their work.

There are 73 species in the Splachnaceae family, and nearly half of these species are dung mosses. Most are found in temperate habitats in both the northern and southern hemispheres, with a few species occurring in the mountains of subtropical regions. They can be found in both wet and relatively dry habitats. Dung mosses are generally fast growing but short lived, with some lasting only about 2 years. It isn’t entirely clear how and why mosses in this family evolved to become entomophilous, but one major benefit of being this way is that their spores are reliably deposited on suitable habitat.

Since Splachnum can grow only on droppings, and nowhere else, the wind cannot be trusted with dispersal. Escape of the spores is successful only if they have both a means of travel and a reserved ticket for a particular destination. In the monotonous green of the bog, flies are attracted to the cotton candy colors of Splachnum, mistaking them for flowers. Rooting about in the moss for non-existent nectar the flies become coated with the sticky spores. When the scent of fresh deer droppings arrives on the breeze, the flies seek it out and leave Splachnum-coated footprints in the steaming dung.

Sporophytes of Splachnum vasculosum (photo credit: www.eol.org)

Sporophytes of Splachnum vasculosum (photo credit: www.eol.org)

References

Koponen, A. 2009. Entomophily in the Splachnaceae. Botanical Journal of the Linnean Society 104: 115-127.

Marino, P., R. Raguso, and B. Goffinet. 2009. The ecology and evolution of fly dispersed dung mosses (Family Splachnaceae): Manipulating insect behavior through odour and visual cues. Symbiosis 47: 61-76.

Year of Pollination: Dung Moss

Last year I wrote about two groups of plants that emit foul odors when they bloom: corpse flowers and carrion flowers. Their scent is akin to the smell of rotting flesh, hence their common names. The purpose of this repugnant act is to attract a specific group of pollinators: flies, carrion beetles, and other insects that are attracted to gross things. Though this particular strategy is rare, these aren’t the only plants that have evolved to produce stinky smells in order to recruit such insects to aid in their reproductive processes. For one, there is a very unique group of mosses that do this, commonly known as dung mosses. Judging from the name, you can probably imagine what their smell must be like. However, their common name doesn’t just describe their scent, but also where they live.

At least three genera (SplachnumTetraplodon, and Tayloria) in the family Splachnaceae include species that go by the common name, dung moss. All Splachnum and Tetraplodon species and many species in the genus Tayloria are entomophilous. Entomophily is a “pollination syndrome”, a subject we will explore more thoroughly in future posts, in which pollen or spores are distributed by insects. Compare this to anemophily, or wind pollination, which is the more common way that moss spores are distributed. In fact, dung mosses are the only mosses known to exhibit entomophily.

Dung Moss (photo credit: wikimedia commons)

Dung Moss (photo credit: wikimedia commons)

Before we go too much further, it’s probably important to have a basic understanding of how mosses differ from other plants. Mosses are in a group of non-vascular and non-flowering plants called bryophytes. Vascular tissues are the means by which water and nutrients are transported to and from different plant parts. Lacking vascular tissues, water and nutrients are simply absorbed by the leaves of bryophytes (although some species have structures akin to vascular tissue), which is why they typically grow low to the ground and in moist environments. Bryophytes also lack true roots and instead have rhizoids, threadlike structures that anchor the plants to the ground or to some other substrate (such as dung).

Another major distinction between bryophytes and other plants is that bryophytes spend most of their life cycle as a haploid gametophyte rather than a diploid sporophyte (haploid meaning that it only has one set of chromosomes; diploid meaning that there are two sets of chromosomes, one from the father and one from the mother). In most plants, the haploid gametophyte is a sperm (pollen) or an egg. In bryophytes, the familiar green, leafy structure is actually the gametophyte. The gametophyte houses sperm and egg cells, and when the egg is fertilized by sperm it forms a zygote that develops into the sporophyte structure which extends above the leafy gametophyte. A capsule at the top of the sporophyte contains spores which are eventually released and, upon finding themselves on a suitable substrate in a hospitable environment, germinate to produce new plants. The spore then is comparable to a seed in vascular, seed-bearing plants.

photo credit: wikimedia commons

photo credit: wikimedia commons

As stated earlier, the spores of most mosses are distributed by wind. Dung mosses, on the other hand, employ flies in the distribution of their spores. They attract the flies by emitting scents that only flies can love from an area on the capsule of the sporophyte called the apophysis. This area is often enlarged and brightly colored in yellow, magenta, or red, giving it a flower-like appearance which acts as a visual attractant. The smells emitted vary depending on the type of substrate a particular species of dung moss has become adapted to living on. Some dung mosses grow on the dung of herbivores and others on the dung of carnivores. Some even prefer the dung of a particular group of animals; for example, a population of Tetraplodon fuegiensis was found to be restricted to the feces and remains of foxes. However, dung is not the only material that dung mosses call home.  Certain species grow on carrion, skeletal remains, or antlers. The smells these species produce attract flies that prefer dead flesh and bone in various states of decay.

Yellow Moosedung Moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit: www.eol.org)

Yellow Moosedung Moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit: www.eol.org)

The spores of dung mosses are small and sticky. When a fly visits these plants, the spores adhere to its body in clumps. The fly then moves on to its substrate of choice to lay its eggs, and the spores are deposited where they will then germinate and grow into new moss plants. Flies that visit dung mosses receive nothing in return for doing so, but instead are simply “tricked” into disseminating the propagules. The story is similar with corpse flowers and carrion flowers; flies are drawn in by the smells and recruited to transmit pollen but receive no nectar reward for their work.

There are 73 species in the Splachnaceae family, and nearly half of these species are dung mosses. These mosses are mostly found in temperate habitats in both the northern and southern hemispheres, with a few species occurring in the mountains of subtropical regions. They can be found in both wet and relatively dry habitats. Dung mosses are generally fast growing but short lived, with some lasting only about 2 years. It isn’t entirely clear how and why mosses in this family evolved to become entomophilous, but one major benefit of being this way is that their spores are reliably deposited on suitable habitat. Because of this directed dispersal, they can produce fewer and smaller spores, which is an economical use of resources.

Sporophytes of Splachnum vasculosum (photo credit: www.eol.org)

Sporophytes of Splachnum vasculosum (photo credit: www.eol.org)

References

Koponen, A. 2009. Entomophily in the Splachnaceae. Botanical Journal of the Linnean Society 104: 115-127.

Marino, P., R. Raguso, and B. Goffinet. 2009. The ecology and evolution of fly dispersed dung mosses (Family Splachnaceae): Manipulating insect behavior through odour and visual cues. Symbiosis 47: 61-76.