The Cypress-Knee Sedge

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Sedges (Carex spp.) simply do not get the attention they deserve. I am part of this problem because like so many others, I have breezed over them in vegetation surveys as “just another graminoid.” This is truly a shame because not only are sedges absolutely fascinating organisms, they are immensely important ecologically as well. I am working hard to get to know sedges better so that I too can fully appreciate their place in our ecosystems. One of the coolest specialist sedges I just recently learned about is the so-called cypress-knee sedge (Carex decomposita). For all intents and purposes, this sedge is considered something of an epiphyte!

The cypress-knee sedge has a fondness for growing on wood. Most often you will find it rooted to the buttresses and knees of bald cypress (Taxodium distichum) or the swollen trunk of a swamp tupelo (Nyssa aquatica). It can also be found growing out of rotting logs that float on the surface of the water. It is a long lived species, with individuals having records stretching back through decades of wetland plant surveys. When supplied with the conditions it likes, populations can thrive. That is not to say that it does well everywhere. In fact, it has declined quite a bit throughout its range.

Juvenile cypress-knee sedges establishing in moss along the water line of a bald cypress.

Juvenile cypress-knee sedges establishing in moss along the water line of a bald cypress.

One of the key wetland features that the cypress-knee sedge needs to survive and prosper is a stable water level. If water levels change too much, entire populations can be wiped out either by drowning or desiccation. Even before the sedge gets established, its seeds require stable water levels to even get to suitable germination sites. Each achene (fruit) comes complete with a tiny, corky area at its tip that allows the seeds to float. Floating seeds are how this species gets around. With any luck, some seeds will end up at the base of a tree or on a floating log where they can germinate and grow. If water levels fluctuate too much, the seeds simply can’t reach such locations.

Its dependence on high quality wetlands is one of the major reasons why the cypress-knee sedge has declined so much in recent decades. Aside from outright destruction of wetlands, changes in wetland hydrology can have dire consequences for its survival. One of the major issues for the cypress-knee sedge is boat traffic. Boat wakes create a lot of disturbance in the water that can literally scour away entire populations from the base of trees and logs. Another major threat are changes to upstream habitats. Any alteration to the watersheds of wetland habitats can spell disaster for the cypress-knee sedge. Alterations to creeks, streams, and rivers, as well as changes in ground water infiltration rates can severely alter the water levels in the swamps that this sedge depends on for survival.

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Closeups of the infructescence showing details of the perigynia (fruit).

Closeups of the infructescence showing details of the perigynia (fruit).

Less obvious threats also include changes in plant cover. If the wetlands in which it grows become too dense, the cypress-knee sedge quickly gets out-competed. To thrive, the cypress-knee sedge needs slightly more sunlight than a densely forested wetland can provide. In fact, some have even noted that cypress-knee sedge populations can explode after selective logging of such wetlands. Such explosions have been attributed to not only extra sunlight but also the addition of woody debris, which provides much needed germination sites. That being said, such explosions can only be maintained if woody debris is left in place and further wetland disturbances do not continue.

The plight of the cypress-knee sedge stands as a reminder of just how poorly we treat wetlands around the globe. Aside from providing valuable ecosystem services for the human environment (flood control, water filtration, etc.), wetlands are home to countless unique species. Only by treating wetlands betters and attempting to restore some of what has been lost will we ever do better by wetland species like the cypress-knee sedge. Hopefully by showcasing species like this, people will begin to feel a little more compassion towards the ecosystems on which they depend. Please consider supporting a wetland conservation and restoration initiative in your region!

Photo Credits : LDWF Natural Heritage Program [1] & Paul Marcum (Midwest Graminoides) [2] [3] [4]

Further Reading: [1] [2]


Botanical Buoys

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American featherfoil (Hottonia inflata) is a fascinating aquatic plant. It can be found in wetlands ranging from the coastal plains of Texas all the way up into Maine. Though widespread, American featherfoil is by no means common. Today I would like to introduce you to this gorgeous member of the primrose family (Primulaceae).

American featherfoil may look like a floating plant but it is not. It roots itself firmly into the soil and spends much of its early days as a vegetative stem covered in wonderful feathery leaves. It may be hard to find during this period as no part of it sticks above the water. To find it, one must look in shallow waters of ponds, ditches, and swamps that have not experienced too much disturbance. More on this in a bit.

American featherfoil lives life in the fast lane. It is what we call a winter annual. Seeds germinate in the fall and by late October, juveniles can be seen sporting a few leaves. There it will remains throughout the winter months until early spring when warming waters signal the growth phase. Such growth is rapid. So rapid, in fact, that by mid to late April, plants are beginning to flower. To successfully reproduce, however, American featherfoil must get its flowers above water.

The need to flower out of water is exactly why this plant looks like it is free floating. The flower stalks certainly do float and they do so via specialized stems, hence the specific epithet “inflata.” Each plant grows a series of large, spongy flowering stalks that are filled with air. This helps buoy the stems up above the water line. It does not float about very much as its stem and roots still anchor it firmly into place. Each inflorescence consists of a series of whorled umbels that vary in color from white to yellow, and even violet. Following pollination, seeds are released into the water where they settle into the mud and await the coming fall.

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As I mentioned above, American featherfoil appreciates wetland habitats that haven’t experienced too much disturbance. Thanks to our wanton disregard for wetlands over the last century or so, American featherfoil (along with countless other species) has seen a decline in numbers. One of the biggest hits to this species came from the trapping of beavers. It turns out, beaver ponds offer some of the most ideal conditions for American featherfoil growth. Beaver ponds are relatively shallow and the water level does not change drastically from month to month.

Historically unsustainable levels of beaver trapping coupled with dam destruction, wetland draining, and agricultural runoff has removed so much suitable habitat and with it American featherfoil as well as numerous wetland constituents. Without habitat, species cannot persist. Because of this, American featherfoil has been placed on state threatened and endangered lists throughout the entirety of its range. With the return of the beaver to much of its former range, there is hope that at least some of the habitat will again be ready for American featherfoil. Still, our relationship with wetlands remains tenuous at best and until we do more to protect and restore such important ecosystems, species like American featherfoil will continue to suffer. This is why you must support wetland protection and restoration in your region!

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

Further Reading: [1] [2]

 

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 Parrot Pitcher Plant

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Southeastern North America is the true home of the carnivorous plants belonging to the genus Sarracenia. Seven of the approximately eight species in this genus reside in North America's coastal plain forests and nowhere else. These evolutionary marvels are famous the world over for their carnivorous pitfall traps but not all members conform to this style of prey capture. The most aberrant of these carnivores is the so-called parrot pitcher plant (Sarracenia psittacina).

The parrot pitcher plant would be easy to pick out of a lineup, even with an untrained eye. Instead of tall, lanky, upright pitchers, it produces a rosette of smaller, entirely prostrate pitchers. Additionally, the leaf-like hood that covers the pitchers of its relatives appears to have grown into a dome-like structure speckled with translucent patches. Finally, the belly of each pitcher sports a leafy fin called an "ala" that runs the whole length of the tube. Indeed, with the exception of perhaps the purple pitcher plant (S. purpurea), the parrot is truly an oddball.

Its unique appearance is likely an adaptation to seasonal flooding and has changed the way in which this particular species captures prey. The pitchers of the parrot pitcher plant do not function as pitfall traps like those of its relatives. Instead, this species utilizes the "lobster trap" method of prey capture. Lured to its pitchers by their bright colors, insects gradually explore the traps. The fin-like ala directs these unsuspecting victims to the mouth of the pitcher. The translucent patches on the domed hood lure the insect into a false sense of security.

Once inside, the insects become disoriented and cannot easily find the proper escape rout. As they crawl farther into the pitcher, backward pointing hairs ensure that escape is impossible. Death is followed by digestion as the pitcher obtains yet another nutrient-rich meal. However, insects aren't the only game in town for the parrot pitcher plant.

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Because of its prostrate habit, the parrot pitcher plant regularly finds itself underwater whenever its already wet habitat floods. This would be bad news for most other pitchers as their upright position would allow whatever was inside to float out and away. Such is not the case for the parrot pitcher. Underwater, the pitchers become even more like a lobster trap. Everything from aquatic insects to tadpoles and fish can and do fall victim to this plant. As such, not even seasonal flooding can put a damper on this unique pitcher plants meal ticket. It is a wonderful example species adaptation.

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Like all members of the southeastern coastal plain community, the parrot pitcher plant is losing its habitat at an alarming rate. Habitat loss is an ever present threat, both in the form of outright destruction from logging and development as well as from sequestration of fire. Coastal plain communities are fire-adapted ecosystems and without it, the myriad species that call this region home are overgrown and choked out. Research has shown that the parrot pitcher plant, as well as other pitcher plants, greatly benefit from regular fires. Fire clears away competing vegetation and the plants respond with vigor.

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Luckily, parrot pitcher plant numbers are stable at this point in time. Its low growth habit saves it from threats like mowing, which means that it can do well in places like roadside ditches that are less favorable for its taller relatives. I have said it before and I will say it again, if you value species like the parrot pitcher plant, please do everything you can to support land conservation efforts. Please check out what organizations such as The Longleaf Alliance, Partnership For Southern Forestland Conservation, The Nature Conservancy, and The National Wildlife Federation are doing to protect this amazing region. Simply click the name of the organization to find out more.

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

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

 

Fish: The Unsung Heroes of Seed Dispersal

Fruits of the tucum palm.

Fruits of the tucum palm.

It goes without saying that effective seed (and spore) dispersal is vital for thriving plant populations. Without it, plant populations will stagnate and disappear. Whereas we know quite a bit about the role animals like birds, bats, and ants play in this process, there is another group of seed dispersers that are proving to be vital to the long-term health and survival of tropical forests around the globe - fish. 

The idea of seed dispersing fish may come as a shock to some but mounting evidence is showing that fruit-eating fish play a major role in the reproductive cycle of many tropical plant species. This is especially true in seasonally flooded tropical forests. To date, more than 100 different fish species have been found with viable seeds in their guts. In fact, some fish species, such as the pacu (Piaractus mesopotamicus), specialize on eating fruits.

A big ol' pacu looking for its next fruit meal.

A big ol' pacu looking for its next fruit meal.

By monitoring how fruit-eating fish like the pacu behave in their environment, scientists are painting a picture of tropical seed dispersal that is quite remarkable. Take, for instance, the tucum palm (Bactris glaucescens). Native to Brazil's Pantanal, this palm produces large, red fruits and everything from peccaries to iguanas will consume them. However, when eaten by these animals, the seed either don't make it through the gut in one piece or they end up being pooped out into areas unsuitable for germination. Only when the seeds have been consumed by the pacu do they end up in the right place in the right condition. It appears that pacus are the main seed dispersal agent for this palm. 

A beautiful tucum palm in the dry season.

A beautiful tucum palm in the dry season.

The tucum palm isn't alone either. The seeds of myriad other plant species known to inhabit such seasonally flooded habitats seem to germinate and grow most effectively only after having been dispersed by fish. Pacus are also responsible for a considerable amount of seed dispersal for plants such as Tocoyena formosa (Rubiaceae), Licania parvifolia (Chrysobalanaceae), and Inga uruguensis (Fabaceae). Even outside of the tropics, fish like the channel catfish (Ictalurus punctatus) are being found to be important seed dispersers of riparian plants such as the eastern swampprivet (Forestiera acuminata).

Camu-camu ( Myrciaria dubia )

Camu-camu (Myrciaria dubia)

Without fish, these plants would have a hard time with seed dispersal in such seasonally flooded habitats. Lacking a dispersal agent, these seeds would be stuck at the bottom of a river, buried in anoxic mud. As fish migrate into flooded forests, they can move seeds remarkable distances from their parents. When the flood waters recede, the seeds find themselves primed and ready to usher in the next generation.

Fruits of the Camu-camu ( Myrciaria dubia ) also benefit from dispersal by fish.

Fruits of the Camu-camu (Myrciaria dubia) also benefit from dispersal by fish.

Not all fish perform this task equally as well. Even within a species, there are differences in the effectiveness of seed dispersal services. Scientists are finding that large fish are most effective at proper seed dispersal. Not only can they consume whole fruits with little to no issue, they are also the fish that are most physically capable of moving large distances. Sadly, humans are seriously disrupting this process in a lot of ways.

For starters, dams and other impediments are cutting off the migratory routs of many fish species. Large fish are no longer able to make it into flooded regions of forest far upstream once a dam is in place. What's more, dams keep large tracts of forest from flooding entirely. As such, fish are no longer able to migrate into these regions, which means less seeds are making it there as well. This is bad news for forest regeneration.

"Gimme fruit" says local channel cat.

"Gimme fruit" says local channel cat.

It's not just dams hurting fish either. Over-fishing is a serious issue in most water ways. Pacus, for instance, have seen precipitous declines throughout the Amazon over the last few decades. Specifically targeted are large fish. Unfortunately, regulations that were put into place in order to help these fish may actually be harming their seed dispersal activities. Fish under a certain size must be released from any catch, thus a disproportionate amount of large fish are being removed from the system.

Logging is taking a serious toll as well. Floodplain forests have been hit especially hard by logging, both legal and illegal. The lower Amazon River, for example, has almost no natural floodplain forests left. Reports from fish markets in these areas have shown fewer and fewer frugivorous fish each year. It would appear that large fruit-eating fish are disappearing in the areas that need seed dispersal the most. It is clear that something drastic needs to happen. At the very least, fruit-eating fish need more recognition for the ecosystem services they provide.

Forest health and management is a holistic endeavor. We cannot think of organisms in isolation. This is why ecological literacy is so important. We are only now starting to realize the role of large fish in forest regeneration and who knows what kinds of discoveries are just over the horizon. This is why land conservation efforts are so important. We must move to protect wild spaces before they are lost for good. Please consider donating to one of the many great land conservancy agencies around the globe. 

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

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

 

Wet Prairies and the White Lady's Slipper

This week we visit a wet prairie in search of the white lady's slipper orchid (Cypripedium candidum). This is a unique habitat type full of incredible plants and we meet many of them along the way. Special thanks to Paul Marcum (http://bit.ly/2r6SG8s) in making this episode possible! 

If you would like to support orchid conservation efforts here in North America, consider purchasing a stick over at http://www.indefenseofplants.com/shop/

Producer, Writer, Creator, Host:
Matt Candeias (http://www.indefenseofplants.com)

Producer, Editor, Camera:
Grant Czadzeck (http://www.grantczadzeck.com)

Twitter: @indfnsofplnts

Facebook: http://www.facebook.com/indefenseofpl...

Patreon: http://www.patreon.com/indefenseofplants

Tumblr: http://www.tumblr.com/indefenseofplants

_________________________________________________________________

Music by: 
Artist: Lazy Legs
Track: Chain of Pink
Album: Chain of Pink
http://lazylegs.bandcamp.com

Thanks, Ducks!

Recent research suggests that certain duck species are crucial for maintaining wetland plant diversity in highly fragmented landscapes. Functioning wetlands are becoming more and more isolated each year. As more land is gobbled up for farming and development, the ability for plants to get their seeds into new habitats is made even more difficult. Luckily, many plants utilize animals for this job. Seeds can become stuck in fur or feathers, and some can even pass through the gut unharmed. What's more, animals can move great distances in a short amount of time. For wetland plants, the daily movements of ducks seems to be paramount. 

By tracking the daily movements of mallards, a team of researchers from Utretch University were able to quantify how crucial these water fowl are for moving seeds around. What they found was quite remarkable. In autumn and winter, the diet of mallards switches over to seeds. Not all seeds that a mallard eats get digested. Many pass through the gut unharmed. Additionally, mallards are strong flyers. On any given day they can travel great distances in search of winter foraging grounds. In the evenings, they return to roosting sites with a high degree of fidelity. 

The research team was able to demonstrate that their movements cover even greater distances in highly fragmented landscapes. It's these daily migrations that are playing a major role in maintaining plant diversity between distant wetlands. This is especially important for wetlands that function as roost sites. Whereas mallards distribute around 7% of the surviving seeds they eat among foraging sites, that number jumps to 34% for surviving seeds at roost sites. Given the sheer number of mallards on the landscape, these estimates can really add up. 

It is likely that without mallards, North American wetlands would be much less diverse given their increasingly isolated nature. However, not all seeds are dispersed equally. Small seeds are far more likely to pass through the gut of a duck unharmed, meaning only a portion of the plant species that grow in these habitats are getting a helping hand (wing?). Still, the importance of these birds cannot be overlooked. The next time you see a mallard, thank it for maintaining wetland plant diversity. 

Photo Credits: [1] [2]

Further Reading: [1]

Why You Should Never Buy Cypress Mulch

Gardening season is soon to be underway here in the northern hemisphere. This past weekend saw droves of people taking advantage of the nice weather by getting their hands dirty in the garden. A walk around the neighborhood brought with it a lot of smiles and a chance to reconnect with neighbors I haven't talked to in a while but it also brought with it something sinister. Lingering in the air was the scent of cypress mulch. Tons upon tons of it are being spread over gardens everywhere. One might ask "Whats the problem? Cypress mulch is more durable and more insect resistant than other mulches!"

WRONG!

Anymore today, these ideas are leftovers of a long gone era. Back when old growth cypress forests were still a thing, these centuries old trees did impart rot and pest resistance into their wood. Today, this is not the case. Because logging has taken most of the old growth cypress from places like Florida and Louisiana, mulch companies have had to resort to cutting down and mulching young, second and third growth cypress stands. Barely given the time to grow into the towering specimens their parents and grandparents once were, these young trees have not yet imparted the centuries worth of compounds into their wood that keep them from rotting and deter insect predators.

The saddest part of the cypress mulch industry is that they are destroying valuable and irreplaceable habitat for the myriad lifeforms that rely on cypress swamps for their existence. To add insult to injury, recovery of cypress trees is almost negligible anymore today due to the way we have managed our waterways. Cypress seedlings require inundation by freshwater and regular silt deposition in order to successfully germinate. A century of flood control, inundation by brackish water, as well as dam and ship canal building have completely upset this dynamic. Now, instead of building new habitat for cypress swamps, these sediments are washed away, far out into the Gulf of Mexico.

What staggeringly few people seem to care to realize is that cypress swamps are our first line of defense against hurricanes. Cypress swamps can cut the force of a storm surge by 90%. It has been estimated that the cypress swamps in Louisiana alone are worth a staggering $6.7 billion in storm protection every year. That is a lot of cash, people!

As with any other industry, the cypress mulch companies are driven by consumer demand. The simple act of individuals, communities, and local governments not purchasing this nasty product is all it will take to lessen the blow to these precious habitats. At the rate cypress is being cut, it will not take long for us to exhaust the resource entirely. As you are looking to do some gardening this year, and many years into the future, please keep these great trees in mind and stop buying cypress mulch. In lieu of wood and bark mulches, you should consider using shredded leaves from your property instead. They make excellent mulch and being locally sourced, the reduce the chances of introducing disease and other pests to your landscape. In the words of Captain Planet, "the power is yours!"

Photo Credit: Jesse Reeder (http://bit.ly/1wmQpn8)

Further Reading: [1] [2] [3] [4]






 

Swamp Pink

The name "swamp pink" just doesn't do Helonias bullata justice. Yes, this species grows in wetlands and yes, the flowers are indeed pink. However, seeing one of these beauties in person will help you realize that the grandeur of such a plant cannot be summed up by any title. Sadly, if we continue to treat wetlands with rampant disregard, future generations will only see swamp pinks in the pages of a book or in an internet photo album.

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As stated, swamp pink likes to have its feet wet. Not just any old wetland will do though. Swamp pinks require a very stable water table with a water line that rests just below the dense rosette of strap-like leaves. At one time, this species could be found from Staten Island, New York all the way south to Georgia. There is even a disjunct population located in the Southern Appalachian Mountains. Today, swamp pink has been reduced to a mere fraction of this former range and now only occurs in isolated pockets of New Jersey, Delaware, Maryland, Virginia, North Carolina, South Carolina, and Georgia.

There are two major threats to the continued survival of this species - wetland destruction and poaching. Because it is a magnificent looking plant, it is often dug up and taken away. Sadly, swamp pink does not transplant well and plants rarely survive the ordeal. Far more deadly to this species is loss of habitat. It isn't just outright destruction of wetlands either. Alterations in the hydrology that stem from increased runoff and poor wetland buffering can cause entire populations to die off.

Both seed production and germination rates are low for this species. What's more, viable seeds suffer from minuscule dispersal distances. Because of this, establishment of new populations can be difficult. Also, since most reproduction is clonal, the gene pools of many extant populations are quite shallow. The plight of the swamp pink really brings meaning to the cultural meme "this is why we can't have nice things."

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

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