Eelgrass Sex is Strange

Pollination may seem like a strange thing to us humans. Whereas we only require two of us to accomplish reproduction, plants have to utilize a third party. The most familiar cases include insects like bees and butterflies. Unique examples include birds, bats, and even lizards. Many plants forego the need of an animal and instead rely on wind to broadcast copious amounts of pollen into the air in hopes that it will randomly bump into a receptive female organ.

This has worked very well for terrestrial plants but what about their aquatic relatives? Water proves to be quite an obstacle for the methods mentioned above. Some species get around this by thrusting their flowers above the surface but others don't bother. One genus in particular has evolved a truly novel way of achieving sexual reproduction without having to leave its aquatic environment in any way.

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Meet the Vallisnerias. Commonly referred to as tape or eelgrasses, this genus of aquatic plants has been made famous the world over by their use in the aquarium trade. In the wild they grow submerged with their long, grass-like leaves dancing up into the water column. Where they are native, eelgrasses function as an important component of aquatic ecology. Everything from fish and crustaceans all the way up to manatees utilize tape grass beds for both food and shelter. Eelgrasses stabilize stream beds and shorelines and even act as water filters.

All this is quite nice but, to me, the most interesting aspect of Vallisneria ecology is their reproductive strategy. Whereas they will reproduce vegetatively by throwing out runners, it is their method of sexual reproduction that boggles the mind. Vallisneria are dioecious, meaning individual plants produce either male or female flowers. The female flowers are borne on long stalks that reach up to the water surface. Once there they stop growing and start waiting. Because of their positioning, water tension causes a slight depression around the flowers at the surface. The depression resembles a little dimple with a tiny white flower in the center.

A female  Vallisneria  flower

A female Vallisneria flower

Male  Vallisneria  flowers floating on the water surface.

Male Vallisneria flowers floating on the water surface.

Male flowers are very different. Much smaller than the female flowers, a single inflorescence can contain thousands of individual male organs. As they mature underwater, the male flowers break off from the inflorescence and float to the surface. Similar to wind pollinated terrestrial plants, Vallisneria use water currents to disperse their pollen. Once at the surface, the tiny male flowers float around like little pollen-filled rafts.

If a male flower floats near the dimple created by a female flower, it will slide down into the funnel-like depression where it will contact with the female flowers. This is how pollination is achieved. Once pollinated, hormonal changes signal the stem of the female flower to begin to coil up like a spring, drawing the developing seeds safely underwater where they will mature. Eventually hundreds of seeds are released into the water currents.

After pollination, the stem of the female flower coils up, drawing the ripening ovaries safely underwater.

After pollination, the stem of the female flower coils up, drawing the ripening ovaries safely underwater.

The Vallisneria are incredible aquatic plants. Their bizarre reproductive strategy has ensured that these plants never really have to leave the water. The fact that they can also reproduce vegetatively means that many species are very successful plants. In fact, some species have become noxious invasive weeds where they have been introduced far outside of their native range. If you own these plants in any way, do take the necessary measures to ensure that they never have the chance to become invasive.

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]

Meet the Crypts

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If you have ever spent time in an aquarium store, you have undoubtedly come across a Cryptocoryne or two. Indeed, these plants are most famous for their indispensable role in aquascaping freshwater aquaria. As organisms, however, crypts receive considerably less attention. Nonetheless, a handful of dedicated botanists have devoted time and effort to understanding this wonderful genus of tropical Aroids. What follows is a brief introduction to the world of Cryptocoryne plants. 

Cryptocoryne is a genus that currently consists of around 60 - 65 species, all of which are native to tropical regions of Asia and New Guinea. Every few years it seems at least one or two new species are added to this list and without a doubt, more species await discovery. All crypts are considered aquatic to one degree or another. Ecologically speaking, however, species fall into four broad categories based on the types of habitats they prefer.

Cryptocoryne cognata in situ .

Cryptocoryne cognata in situ.

The most familiar crypts grow along the banks of slow-moving rivers and streams and find themselves submerged for a large portion of their life. Others grow in seasonally flooded habitats and experience a pronounced dry season. These species usually go dormant until flood waters return. Still others can be found growing in swampy forested habitats, often in acidic peat swamps. Finally, a few crypts have adapted to living in tidal zones in both fresh and brackish waters.

Like all aquatic plants, crypts face a lot of challenges living in water. One of the biggest challenges is reproduction. Despite their aquatic nature, crypts will not flower successfully underwater. If growing submerged, most crypt species reproduce vegetatively via a creeping rhizome. As such, crypts often form large, clonal colonies in both the wild and in aquaria, a fact that has made a few crypts aggressive invaders in places like Florida.

Cryptocoryne wendtii  is one of the most common species in the aquarium trade. Its textured leaves are thought to have a higher surface area, allowing this plant to thrive in shaded aquatic habitats.

Cryptocoryne wendtii is one of the most common species in the aquarium trade. Its textured leaves are thought to have a higher surface area, allowing this plant to thrive in shaded aquatic habitats.

Given proper hydrologic cycles, however, crypts will flower and when they do, it is truly a sight to behold. As is typical of aroids, crypts produce an inflorescence comprised of a spadix with whirls of male and female flowers covered by a decorative sheath called a spathe. This spathe is the key to successful flowering among the various crypt species.

Species like  C. becketti  have become invasive in places like Florida, no doubt thanks to aquarium hobbyists.

Species like C. becketti have become invasive in places like Florida, no doubt thanks to aquarium hobbyists.

If the spathe were to open underwater, the inflorescence would quickly rot. Instead, most crypts seem to have an uncanny ability to sense water levels. At early stages of development, the spathe completely encloses the developing spadix in a water tight package. The tubular spathe continues to grow upward until the top has breached the surface. Consequently, the overall length of a crypt inflorescence is highly variable depending on the water level of its habitat. Crypts living in tidal zones take this a step further. Somehow they are able to time their flowering events to the ebb and flow of the tides, only producing flowers during periods of the month when tides are at their lowest.

Cryptocoryne ligua

Cryptocoryne ligua

With the tip of the inflorescence safely above water, the spathe will finally open revealing their surprisingly complex anatomy and coloration. It is a shame that most crypt growers never get to see such floral splendor in person. The spathe of many crypt species emit a faint but unpleasant odor. Additionally, some species adorn the spathe with fringes that, coupled with stark coloration, is thought to improve the chances of pollinator visitation.

Pollinators are poorly studied among crypts, however, it is thought that small flies take up the bulk of the work. Lured in by the promise of a rotting meal on which they can feed and lay their eggs, the flies become trapped inside the long tube of the spathe. Like the pitfall traps of a pitcher plant, the inner walls of the spathe are coated in a waxy substance that keeps the insects from crawling out before they do their job.

In general, the female flowers mature first. If the insect inside has visited a crypt of the same species the day before, it is likely carrying pollen and thus deposits said pollen onto the stigmas of the current crypt. After the female flowers have had a chance at being fertilized, the male flowers then mature. The insects inside are then dusted with new pollen, the walls of the spathe lose their slippery properties, and the insects are released in hopes of repeated the process again.

The fruit of a  Cryptocoryne  is called a syncarp.

The fruit of a Cryptocoryne is called a syncarp.

To the best of my knowledge, most crypts are not self-compatible. Instead, plants must receive pollen from unrelated individuals to set seed. Because large crypt colonies are often made up of clones of a single mother plant, sexual reproduction can be rather infrequent among the various species. Nonetheless sexual reproduction does occur and the seeds are produced in a different way than most other aroids. Instead of berries, crypts produce their seeds in a aggregated collection of fruits called a syncarp. When ripe, the syncarp opens like a little star and the seeds float away on the current.

One species, Cryptocoryne ciliata, takes seed production to a whole different level by producing viviparous seeds. Before the syncarp even opens, the seeds actually germinate on the mother plant. In this way, tiny seedlings complete with roots and leaves are released instead of seeds. Seedlings have a much greater surface area than seeds and readily get stuck in mud as well as other aquatic vegetation. In this way, C. ciliata offspring get a jump start on the establishment process. It is no wonder then that C. ciliata has one of the widest distributions of any of the crypt species.

Cryptocoryne ciliata

Cryptocoryne ciliata

Despite plenty of overlap among the ranges of various crypt species, the genus displays an amazing array of variation. Some have likened crypts to Araceae's version of Darwin's finches in that the unique ecology of each species appears to have created barriers to species introgression. Though hybrids do occur, each crypt seems to maintain its own niche via a unique habitat requirement, differing flower phenology, or a specific set of pollinators. It would appear that much can be learned about the mechanics of speciation by studying the various Cryptocoryne and their habits.

Unfortunately, the limited geographic distribution and specific habitat requirements of crypt species is cause for concern. Many are growing more and more rare as human settlements expand and destroy valuable crypt habitat. As popular as some crypts may be in cultivation, many others have proven too idiosyncratic to grow on a commercial level. More work is certainly needed to properly assess populations and bring plants into cultivation as a form of ex situ conservation.

Cryptocoryne cordata  Var. Siamensis 'Rosanervig' is a contoversial variety names recognized by the stark patterns of venation on its leaves.

Cryptocoryne cordata Var. Siamensis 'Rosanervig' is a contoversial variety names recognized by the stark patterns of venation on its leaves.

Proper study is further complicated by the fact that many crypt species are highly plastic. They have to be in order to survive the rigors of their aquatic environment. True species identification can really only be assessed when flowers are present and some populations seem to prefer vegetative over sexual reproduction a majority of the time. A multitude of subspecies exist, though the degree to which they should be formally recognized is up for debate.

I think it is safe to say that Cryptocoryne is a genus worth far more attention than it currently receives. They are without a doubt important components of the ecology of their native habitats and humans would do well to understand them a bit better. With a bit more attention from botanical gardens and other conservation organizations, perhaps the future for many crypts does not have to be so bleak.

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

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