Vernal Pools: The Kingdom of the Amphibians

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Ovipositing Spotted Salamander

There are few things more magical, few moments more memorable, and few experiences more rewarding than exploring a vernal pool in late winter and early spring.  I wish everyone could know the joy that I feel standing ankle deep in the frigid water amid a deafening cacophony of Spring Peepers and chorus frogs.

Vernal pools are isolated wetlands that form in upland depressions.  In the great forests of the eastern United States, of which the Pineywoods of East Texas are essentially an extension, they often occur in open, mature forests over pockets of dense clay scattered among a matrix of otherwise loamy soil.  They may also occur in the scars of old stream beds left after some waterway changed course millennia ago.  The unique topography of these areas result in pools that hold water during the winter and spring and dry in the summer and fall.

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A dry vernal pool in fall

Vernal pools begin to fill in the late fall and early winter with the return of regular rainfall patterns associated with frequent fronts.  At the same time trees and other plants are entering a period of dormancy, when their water requirements are reduced dramatically.  This reduction in plant activity results in a rising of the water table.  In more northerly climates the pools may not fill until the spring snow melt funnels water to them.  By later spring and early summer, the water table begins to drop as thirsty roots draw from it.  Evapotranspiration increases, and by late summer these depressions will be dry as a bone.

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A full vernal pool in late winter

The ephemeral nature of these wetlands means that fish are unable to survive here.  This does not mean that vernal pools are devoid of life, however.  A multitude of invertebrates specialize in vernal pools and a spectacular array of amphibians depend on them to breed.  It is, in fact, the absence of fish that allow these organisms to thrive here.  Without these voracious piscene predators, and with an abundance of invertebrate prey, amphibian larvae can flourish.

Many amphibians typical of eastern North America reach the southwestern extend in East Texas.  Most of these species breed during the first substantial warm rains of late winter and early spring, often undergoing mass migrations from upland refugia to reach the vernal pools.  Easily the most spectacular of these amphibians, and perhaps my favorite animal of all time, is the Spotted Salamander (Ambystoma maculatum).  These large, brightly colored members of the family Ambystomatidae are like the “poster children” of vernal pools.  Their fate is so closely tied to these unique habitats that they are indicator species for them.  And as healthy vernal pools depend on a healthy, mature forest, the Spotted Salamander is, in turn, an indicator species for the health of the entire forest.

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A male Spotted Salamander

Spotted salamanders spend the majority of their lives confined to small burrows, generally excavated by some species of rodent or other fossorial creature.  As the warm winter rains begin to saturate the soil they slowly emerge from their underground haunts and make their way, en masse, to the vernal pools of their birth.  The males arrive first.  They can be identified this time of year by their swollen cloacas.  When all is said and done they will outnumber the females several to one.  After entering the water they dutifully set out to find the perfect spot to deposit their spermatophore, a gelatinous deposit containing their genetic material.  Here they await the arrival of the females.

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A Spotted Salamander emerges from the leaf litter

When the females do arrive their are swollen with a gelatinous mass of eggs within.  The males dance and display around their spermatophore with a series of flips, gyrations, and underwater acrobatics.  Once they have caught the eye of a female they gently guide her over their spermatophore.  She picks it up with her cloaca and uses it to fertilize her eggs.  She then deposits an impossibly massive egg mass on a submerged twig or vegetation.  The salamanders do not dawdle in the ponds, and as soon as they arrived they return to their upland burrows.  Here they will remain for the rest of the year.  Part of what makes seeing a Spotted Salamander so special is knowing that they are only visible to the above-ground world for a few shorts weeks of the year.

The salamander eggs are protected by a thick gelatinous mass that is resistant to desiccation should the pool dry for a short time during their development.  The Spotted Salamander have a mutualistic relationship with an algae, Oophila amblystomatis, which is found exclusively within the salamanders’ egg masses.  The algae benefits from the nitrogen-rich waste products produced by the developing embryos while the embryos themselves are oxygenated through the biproducts of the photosynthesizing algae.

Eventually tiny, legless gilled salamander larvae will emerge from the egg masses.  They will feed and grow within the vernal pools until sometime in late spring when their legs and lungs are fulled developed.  They will then absorb their gills, lose their tail fins, and emerge from the water to begin a terrestrial life.

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Spotted Salamander Egg Mass

There are other species of salamander that utilize vernal pools.  The Mole Salamander (Ambystoma talpoideum) is a small, stocky salamander of the southeastern United States.  They occur in a variety of ephemeral wetlands, from vernal pools to permanent fish-free ponds within longleaf pine savannahs.  Populations in East Texas often contain both terrestrial and neotenic adults.  In neoteny, adults retain juvnile characteristics; in this case gills and aquatic morphology.  These neotonic adults enjoy an entirely aquatic existence, even breeding in the water.  If the pool begins to drain, however, these aquatic adults can absorb their gills and morph into terrestrial adults.

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Mole Salamander

In East Texas, Smallmouth Salamanders (Ambystoma texanum) seem to be most common in large ephemeral wetland complexes in floodplain forests.  They will still utilize traditional vernal pools, however.  They are variable in pattern, having a variety of lichen-like blotches and flecks ranging from blue-gray to rusty brown.  In Texas they occur in scattered populations throughout much of the eastern third of the state.

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Smallmouth Salamander

The Marbled Salamander (Ambystoma opacum) is an anomaly.  While the other Ambystomatid salamanders in East Texas breed in the late winter/early spring, the Marbled Salamander breeds in the fall, when the vernal pools are still dry.  During autumn’s first cool rains they migrate to the dry pool basins.  Here they breed on the land, and the female deposits her eggs on dry earth beneath logs and other natural debris within the depression.  Here she will stay, guarding her eggs, until the winter rains fill the pools and inundate the eggs.

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A female Marbled Salamander guarding her eggs.

There is method to this madness, however.  By the time the eggs of the Spotted, Mole, and Smallmouth Salamanders have hatched, the Marbled Salamander larvae have been growing for weeks.  What ensues is a mass slaughter of intra-generic predation, as Marbled Salamander larvae consume hundreds of their congenerics.  This spring we observed hundreds of Spotted Salamander egg masses, and most had at least one Marbled Salamander larva sitting atop them.  For a short while at least, the Marbled Salamander is the king of the vernal pool.

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A male Marbled Salamander

Marbled Salamander adults are sexually dimorphic.  Males have bright white to silvery dorsal markings while the markings of the female are a duller silver or copper.  Males arrive to the dry pools first, and as with the Spotted Salamander, significantly outnumber the females.

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Marbled Salamander male (upper right) and female (lower left)

The above mentioned salamanders all have fascinating life histories, but they pale in comparison to the complexity of the life cycle of the Central Newt (Nothophthalmus viridescens louisianensis).  The Central Newt starts life as an aquatic larva emerging from eggs laid in the water.  After a short time they morph into a terrestrial juvenile known as an “eft”.  During this time their skin is noticeably dry to the touch.  After spending a couple of years on land, these juveniles typically return to the water, where they will morph again into an aquatic adult.

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Central Newt Eft

To those exploring a vernal pool at night, their eyes are undoubtedly drawn to the salamanders swimming below the water’s surface.  Their ears, however, belong to the frogs.  Depending on the time of year, a multitude of intricate frog calls may ring out, often in deafening spectacularity.  One of the best known of these anurans is the Spring Peeper (Pseudacris crucifer).  These tiny harbingers of spring fill the night air with their peeping calls that seem too loud to come from such a tiny thing.

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Spring Peeper

Our East Texas representative of the Pseudacris feriarum complex is the Cajun Chorus Frog (Pseudacris fouquettei).  These diminutive frogs seem to breed just about anywhere that water remains for more than a few weeks.  Their call has been likened to a finger running through the teeth of a comb.  In my area P. fouquettei typically starts calling in early to mid December.  I have found many a good vernal pool by following the calls of the Cajun Chorus Frog.

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Cajun Chorus Frog

The Gray Tree Frogs (Hyla chrysoscelis and Hyla versicolor) typically begin breeding in mid Spring, later than the Spring Peeper and Cajun Chorus Frog.  The two species of Gray Tree Frog can only be differentiated by their calls.  In East Texas they typically begin calling in late February/Early March, and may call well into the summer following heavy rains.

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Cope’s Gray Tree Frog (Hyla chrysoscelis)

Perhaps my favorite anuran inhabitant of vernal pools is the Pickerel Frog (Lithobates [Ranapalustris).  They can be differentiated from the similar Southern Leopard Frog by the two rows of large-block like blotches on their back and bright yellow inner legs.  While they are common throughout much of their range, the Pickerel Frog is uncommon in East Texas.  Along with the Southern Leopard Frog, it is one of the earliest breeding Ranids in the Pineywoods.  The Pickerel Frog is known for its toxicity.  Some people who handle these frogs report sensations from mild tingling to moderate pain in their fingers afterwards.  Fortunately I have never had a reaction.  They are also toxic to other amphibians, and may kill other species that are confined to plastic bags with them.

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Pickerel Frog

The Southern Leopard Frog (Lithobates sphenocephalus [Rana sphenocephala]) is one of our commonest Ranids.  I remember the first time I heard their haunting call.  I thought I was listening to some dispute between raccoons or some similar mammal.  Southern Leopard Frogs are quite variable, but their spots are always smaller and more randomly distributed than those of the Pickerel Frog.  They also lack the bright yellow inner legs of their toxic cousins.

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Southern Leopard Frog

The annual amphibian migration to vernal pools is truly one of nature’s great spectacles.  I remember one night a few years ago when Carolina and I, accompanied by some of our closest friends, went out to a vernal pool at night.  A warm front had moved in, drenching East Texas with bands of rain throughout the day and into the night.  As we began travelling to our destination the rain began to pick up, and branched lightning descended from the clouds, dancing across the sky and illuminating the columns of shadow in the dark forest. I was full of anticipation and doubt as we approached the pond. Though the conditions seemed perfect, I had tried this very thing before and turned up empty handed.

Almost inexplicably, at nearly the exact moment we arrived at the pond, the rain had stopped. I rushed to the ponds edge, flashlight in hand. Immediately it became clear that all of my doubts were for naught. As my light crossed the leafy bottom of the vernal pool I could see hundreds of yellow spots; Spots decorating the backs of dozens of Spotted Salamanders that darted back and forth beneath the clear water. We did it, I thought, we timed it perfectly. The calm night was interrupted by the voices of innumerable Cajun chorus frogs that sang from the pool’s edge. It was, without a doubt, one of the most incredible natural events I had ever witnessed.

It was at that moment, when I thought that things couldn’t be more perfect, that my heart could not be more whole, that Carolina said, “look at the sky.” I looked up and saw that the clouds had broken, and in their wake the brilliance of millions of stars shone in the moonless night sky. And though I may be a secular person, the sensation I felt at that moment was no less profound than some deeply religious experience. For the most incredible moments in life need not be some miracle brought on by divine powers, but rather the countless natural wonders of the very world to which we all are a part.

Fall into Winter: November and December Recap

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Fall color along a forest stream

We’re in the heat of winter here in the Pineywoods, and I’ve got a backlog of posts to catch up on.  Soapwort Gentian ended up being the last species checked off my list in 2017.  Though I would not see any more of my “target species”, my November and December were still filled with incredible biodiversity and natural beauty.

In mid November Carolina and I met up with our friend Skip Pudney in the Big Thicket.  We were hoping to photograph the rare orchid Spiranthes longilabris in bloom.  While we did find a single plant, the true show was put on by the invertebrates – pollinators taking advantage in a flush in late season flowers.  We noticed several Yellowjacket Hover Flies (Milesia virginiensis), Ornate Bell Moths (Utetheisa ornatrix), Common Buckeyes (Junonia coenia), Blister Beetles (Epicauta sp.) and more.

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Yellowjacket Hover Fly

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Ornate Bell Moth

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Common Buckeye

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Blister Beetle

By mid-November the leaves had begun to change.  This was a good year for fall color in East Texas.  The leaves of deciduous trees begun to change color in the falls when the days are sunny and the evenings are crisp.  These cues, along with the shortening photoperiod trigger a chemical reaction within the leaves.  Production of chlorophyll halts, and slowly this green pigment begins to break down and is rendered clear.  As the green fades, other colors such as carotenoids and anthocyanins, which have been active in the leaves all along, now become dominant, and the forest turns from green to brilliant hues of yellow, orange, and red.

In late November I spent a foggy morning photographing the maples, oaks, hickories, and elms of a rich hardwood stream bottom.

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Fall color in the fog

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Hickories and maples provide the yellows in this fall forest

I then went on to a steep bluff over the upper reaches of the Neches River, where the Red Maples lived up to their name.

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Fall color on a bluff over the Neches River

While I find broad views of a fall forest to be especially beautiful, the subtle beauty of fall can be observed up close, like in the leaves of poison ivy, as seen below…

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Fall color in poison ivy

…and in the layers of Florida Maple leaves on branches draped around the trunks of pine trees.

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Florida maples display their fall colors.

In early December much of East Texas was hit with an uncharacteristic snow storm.  In over 20 years in the region, I have only seen snow a handful of times, and of those only a fraction actually stuck.  This was one of the finest in recent memories.  In East Texas fall color lingers well into December, and the result was what looked to be a battle between fire and ice.

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Fire and Ice

Deeper in the forest, the landscape appeared a winter wonderland.

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Pineywoods Snowscape

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My Winter Wonderland

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Snow in the Beechwoods

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Dressed in White

Having grown up in Chicago, I experienced harsh, snow-filled winters in my childhood.  It was good to spend some time walking and playing in the snow again – like reuniting with an old friend.  I think that for Caro it was even more special, as she seldom saw snow in her native province of Entre Rios, Argentina.

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Carolina in the Snow

Later in December we visited a longleaf pine savannah shortly after a prescribed fire.  Here we saw fresh cones on the torched leaf litter…

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Borne of Ashes

…and a freshly germinated seedling rising from the ashes. With luck, this tiny seedling will grow into a stately tree in this longleaf pine savannah. Perhaps it will one day harbor the cavity of a Red-cockaded Woodpecker, and Bachman’s Sparrows and Brown-headed Nuthatches will sing from its boughs while Louisiana Pine Snakes and Wild Turkey patrol among its roots.

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New Beginnings

Within the longleaf pine savannah we found Riddell’s Spike-Moss (Selaginella corallina) growing in the crevices of exposed boulders of the Catahoula Formation.  S. corallina is a primitive vascular plant that is typically included with the “fern-allies”, and despite its name is more closely related to ferns than mosses. It has an interesting disjunct range, with one population in central and east Texas, northwestern Louisiana, western Arkansas and Eastern Oklahoma, and another in Alabama and Georgia. I seldom encounter them in East Texas. When I do, it is generally growing off the faces of sandstone outcrops or in areas of deep sand.

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Riddell’s Spike Moss

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Ridell’s Spike Moss

After exploring the savannah we ventured to the bluffs along the Neches River.  Here we found patches of fall color lingering in the American Beech trees on the bluffs’ slopes.  American Beech is one of the last trees to turn in the fall, and they brighten the otherwise gray December forest.  In the photo below the Neches River is visible in the distance.

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The Bluff’s Edge

In late December our friend Scott Wahlberg, Carolina, and I spent the day scouting salamander locations for the spring.  Though the conditions for finding a salamander weren’t ideal, we did turn up a single, apparently gravid, female Spotted Salamander (Ambystoma maculatum).

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Spotted Salamander

Toward the end of December James and I spent some time looking for birds in a local park.  There I was able to capture an image of a Swamp Sparrow (Melospiza georgiana) in the dense underbrush along the margins of a pond using James’s new 600mm lens (more on that in the future).  These striking sparrows are “skulkers” – small birds that prefer dense cover.  I was lucky to get a shot of one as it momentarily paused in its dense domain of tangles.

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Swamp Sparrow

I was also able to photograph a Yellow-rumped Warbler (Setophaga coronata) there.  The birds that winter in East Texas are members of the “myrtle” race, so named because they are one of the few birds that will regularly eat wax-myrtle berries.

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Yellow-rumped Warbler

On December 30, Carolina and I found a Broad-banded Water Snake (Nerodia fasciata confluens) while exploring the bottomlands off the Neches River.  The temperatures hovered just above freezing, and the snake could barely move, yet it was alive and well.  After taking a few photos we left it to weather the cold, as it and its kind have done for countless generations.

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Slow and Steady

The first month of 2017 has passed by and I have yet to cross any species off my list.  Most of the species I put on the list are seasonal, so I spent January focusing on the few resident species that I included, namely the North American River Otter (Lontra canadensis).

Recently my good friend James Childress found a population close to home.  My wife Carolina and i began visiting the site on the weekends, sometimes 2-3 times per day to no avail.  However the waterway and adjacent wetlands and pine-oak uplands are rich with wildlife, and I found many photographic subjects in the otters’ absence

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Wood Duck Drake

.One group that is noticeably missing from my list are birds.  I didn’t include them because though I love bird photography, it is a difficult endeavor, and results are often unpredictable.  If I were to have made a list, however, I would have incorporated the Wood Duck (Aix sponsa).  These denizens of creeks, backwater sloughs, flooded oxbows, and ephemeral ponds can be abundant in parts of The Pineywoods.  Unfortunately they are wary and secretive, and typically all I have to show from encountering one is a memory of splashing, their whistle like vocalization as they retreat, and if I’m lucky a distant glimpse of them flying through the trees.  While pursuing the otters, however, I found a less wary subject and with a bit of patience and luck I was able to finally capture some images of what many consider to be North America’s most beautiful duck.

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Wood Duck Drake

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Wood Duck Drake

One afternoon, as I was trekking with my photographic gear to a spot where we expected the otters might show themselves, I heard my wife call out frantically.  Not in fear, but rather a sense of urgency that I might miss the treasure she had stumbled upon.  She had found a large Regal Jumping Spider (Phidippus regius).  I had tried numerous times to photographic this species, but found them too wary and difficult to approach.  But on this warm January day I found a bold, curious specimen that made for a perfect photographic subject.

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Regal Jumping Spider

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Regal Jumping Spider

I have been photographing the Pineywoods of East Texas for many years now.  While this species list project is a way to add a little spice to my photographic pursuits, I still love wandering the woods with a camera in hand, and no particular goal in mind.  On New Year’s Day Carolina and I went to one of our favorite spots.  We wandered the longleaf pine forest, admiring the fungi taking advantage of the moisture and humidity that remained after some recent rains.  As is typical in our forest wanderings, Carolina had the best find of the day – A pair of Indian Pipes (Monotropa uniflora) pushing skyward from the dense carpet of longleaf pine needles.  I was surprised to see it blooming so late in the season.  The habitat also seemed unusual to me, as I was accustomed to seeing it in the rich loams of moist slopes and stream bottoms.  Pleased with the days observations, we sat among the longleaf pines as the day faded, and contemplated the adventures to come.

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The Fly Agaric (Amanita muscaria) is showy, highly toxic mushroom that appears in pine-dominated forests following late fall and winter rains.

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Ravenel’s Stinkhorn (Phallus ravenelii)

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Unlike most plants, which obtain energy through photosynthesis, Indian Pipe is a mycoheterotroph.  It obtains its energy and nutrients from the mycorrhizal fungi of tree roots.  I have read that it is named for an old Cherokee legend that states that these plants grow where old chiefs are buried so they may have something to smoke in the afterlife.

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Longleaf Pine Savannah

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An old, gnarled post oak growing among longleaf pines in an upland savannah.

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Merlin

Opting for a change of scenery, we decided to take  our pursuit of the North American River Otter to the coast.  I used to see them regularly in the wetlands around Anahuac National Wildlife Refuge when I worked there, and had heard of some recent sightings.  Hoping that we might get lucky Carolina, my mom and I spent a day exploring the Upper Texas Coast.  Though our mammalian quarry eluded us, we did see a number of birds including a variety of wintering waterfowl and wading birds.  The highlight of the day was toss up between a cooperative male Merlin ((Falco columbarius) that perched on a rustic fence post in perfect light, and an American Bittern (Botaurus lentiginosus) that foraged in a ditch next to the road.  Relying on its camouflage it allowed us to approach to within feet of it, as it slowly stalked the reeds.

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American Bittern

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Ringneck Duck (Aythya collaris)

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Spotted Salamanders en route to a vernal pool.

Back in the Pineywoods I checked the weather daily, awaiting the conditions that would bring about one of nature’s great events – the Spotted Salamander migration.  Among the world’s most spectacular amphibians, the Spotted Salamander (Ambystoma maculatum) makes its home in the forests of the eastern United States.  Spotted Salamanders spend the majority of the year underground, hidden from the world in small burrows that have been excavated by rodents or other species.  In response to warm rains in later winter/early spring they emerge en masse and migrate to their breeding ponds, where they form large breeding congresses in order to propagate the next generation.  Finally we had a night with the perfect conditions and Carolina and I visited some breeding sites with our friends James and Erin Childress, where we observed hundreds of Spotted Salamanders and a handful of Mole Salamanders (Ambystoma talpoideum) swimming about the vernal pools.

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A Spotted Salamander male awaits the females’ arrival to his breeding pool.

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A gravid female Spotted Salamander

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A gravid female Mole Salamander

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A false bottomland in the East Texas Pineywoods.  False bottomlands are unique communities that occur in clay-bottomed dpressions located within upland communities.  They typically flood during the winter and spring, and become dry during the summer.  Though not associated with waterways, the community is somewhat similar to bottomland hardwood forests.  Dominant species include willow oak, overcup oak, and mayhaw.  The ephemeral nature of flooding in these communities makes for excellent amphibian habitat.

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An unidentified fungus growing on a fallen tree.

The salamanders were a nice distraction, but it was soon time to return to my pursuit of the river otter.  For Christmas my wife got a remote game camera.  Following the news that James had seen otters, she began setting the camera in areas where we observed ample otter sign.  For a couple weeks she came up empty handed.  One day, while en route to set up the cameras I caught a glimpse of something moving across the leaf litter.  It turned out to be a Red-bellied Snake (Storeria occupitomaculata), a small snake of the eastern U.S. that reaches the southwestern extent of its range in East Texas.  Though common throughout most of its range, the Red-bellied Snake is quite rare in Texas, and I took our encounter to be a good sign for things to come.

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Florida Red-bellied Snake (Storeria occipitomaculata obscura)

Carolina chose a place to set the camera adjacent to some logs that were covered in otter scat and hoped for the best.  Sure enough, the next time we checked the camera’s accumulated images, among the hundreds of photos of branches blowing in the wind, songbirds, and rats, we caught sight of our objective – a North American River Otter.

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With renewed energy I will continue to pursue the aquatic mammal in hopes of capturing its photograph, because slow and steady wins the race.