It may look like just a puddle in the woods, but this is a very specialized wetland habitat, a habitat that is quickly disappearing from the Lower Susquehanna River Watershed. It’s a vernal pool—also known as a vernal pond or an ephemeral (lasting a short time) pool or pond.
Viable vernal pools have several traits in common…
They contain water in the spring (hence the name vernal).
They have no permanent inflow or outflow of water.
They typically dry up during part of the year—usually in late summer.
They are fish-free.
They provide breeding habitat for certain indicator species of forest-dwelling amphibians and other animals.
They are surrounded by forest habitat that supports the amphibians and other vernal pool species during the terrestrial portion of their life cycle.
To have a closer look at what is presently living in this “black leaf” vernal pool, we’re calling on the crew of the S. S. Haldeman to go down under and investigate.
Along the surface of the pool we’re seeing clusters of amphibian eggs, a sign that this pond has been visited by breeding adult frogs and/or salamanders during recent weeks.Amphibian eggs and the white tail filaments of an invertebrate of interest, Springtime Fairy Shrimp (Eubranchipus vernalis), an endemic of vernal ponds.
Let’s take it down for a better look. Dive, all dive!
Algae provides food for the shrimp and other inhabitants of the pool. Leaf litter furnishes hiding places for the pool’s many inhabitants.These loose clusters of eggs appears to be those of Wood Frogs, a vernal pool indicator species.Clusters of Wood Frog eggs, the embryos within those in the center of the image less developed than those to the left.More Wood Frog eggs. Hatching can take anywhere from two weeks to two months, depending on temperature.Wood Frog eggs with developing larvae (tadpoles) plainly visible. The green color of the eggs is created by a symbiotic algae, Oophila amblystomatis, a species unique to vernal pools. The algae utilizes the waste produced by the developing embryos to fuel its growth and in return releases oxygen into the water during photosynthesis. Upon hatching, the tadpoles rely upon the algae as one of their principle food sources.A zoomed-in view showing development of the larvae and what appears to be a tiny invertebrate clinging on the white egg in the upper right. White eggs don’t hatch and may be infected by a fungus.Wood Frog eggs and Springtime Fairy Shrimp.Wood Frog eggs and Springtime Fairy Shrimp.Springtime Fairy Shrimp swim upside-down. Note the small, bluish clusters of eggs attached to the abdomens of these females. Springtime Fairy Shrimp live their entire lives in the vernal pool. After being deposited in the debris at the bottom of the pool, the eggs will dry out during the summer, then freeze and re-hydrate before hatching during the late winter.A damselfly larva consuming fairy shrimp. (Visible in the margin between the uppermost lobes of the dark-colored oak leaf to the right.)Getting in close we see A) the damselfly larva eating a Springtime Fairy Shrimp and B) one of several discarded exoskeletons of consumed shrimp near this predator.A fishfly larva (Chauliodinae). Mosquito numbers are kept in check by the abundance of predators in these pools.Springtime Fairy Shrimp and a Marbled Salamander (Ambystoma opacum) larva. The presence of these species confirms this small body of water is a fully-functioning vernal pool.Springtime Fairy Shrimp and two more larval Marbled Salamanders. The salamanders’ enlarged gills are necessary to extract sufficient oxygen from the still waters of the pool.The Marbled Salamander is one of three species of mole salamanders found in the Lower Susquehanna River Watershed. All breed in vernal pools and live their air-breathing adult lives under the leaves of the forest in subterranean tunnels where they feed on worms and other invertebrates. Photos of an adult Marbled Salamander and the other two species, Spotted Salamander (Ambystoma maculatum) and Jefferson Salamander (Ambystoma jeffersonianum), can be found by clicking the “Amphibians” tab at the top of this page.Marbled Salamanders lay their eggs during the fall. If the bed of the pool is dry at breeding time, the adult female will remain to guard the eggs until rain floods the pool. The eggs hatch upon inundation, sometimes during the winter.Marbled Salamanders, like all amphibians that develop in vernal pools, must complete transformation into their air-breathing terrestrial life stage before the pool dries up in the summer heat.A larval Marbled Salamander explores the bottom of the pool.A larval Marbled Salamander, Wood Frog eggs, and Springtime Fairy Shrimp, it’s an abundance of life in what at first glance may appear to be just a mud puddle.
We hope you enjoyed this quick look at life in a vernal pool. While the crew of the S. S. Haldeman decontaminates the vessel (we always scrub and disinfect the ship before moving between bodies of water) and prepares for its next voyage, you can learn more about vernal pools and the forest ecosystems of which they are such a vital component. Be sure to check out…
If you are a landowner or a land manager, you can find materials specifically providing guidance for protecting, restoring, and re-establishing vernal pool habitats at…
Within the last few years, the early-summer emergence of vast waves of mayflies has caused great consternation among residents of riverside towns and motorists who cross the bridges over the lower Susquehanna. Fishermen and others who frequent the river are familiar with the phenomenon. Mayflies rise from their benthic environs where they live for a year or more as an aquatic larval stage (nymph) to take flight as a short-lived adult (imago), having just one night to complete the business of mating before perishing by the following afternoon.
In 2015, an emergence on a massive scale prompted the temporary closure of the mile-long Columbia-Wrightsville bridge while a blizzard-like flight of huge mayflies reduced visibility and caused road conditions to deteriorate to the point of causing accidents. The slimy smelly bodies of dead mayflies, probably millions of them, were removed like snow from the normally busy Lincoln Highway. Since then, to prevent attraction of the breeding insects, lights on the bridge have been shut down from about mid-June through mid-July to cover the ten to fourteen day peak of the flight period of Hexagenia bilineata, sometimes known as the Great Brown Drake, the species that swarms the bridge.
An adult (imago) male Great Brown Drake (Hexagenia bilineata) burrowing mayfly. Adult mayflies are also known as spinners.A sub-adult (based on the translucence of the wings) female burrowing mayfly (Hexagenia species). The sub-adult (subimago or dun) stage lasts less than a day. Normally within 18 hours of leaving the water and beginning flight, it will molt into an adult, ready to breed during its final night of life.
After so many years, why did the swarms of these mayflies suddenly produce the enormous concentrations seen on this particular bridge across the lower Susquehanna? Let’s have a look.
Following the 2015 flight, conservation organizations were quick to point out that the enormous numbers of mayflies were a positive thing—an indicator that the waters of the river were getting cleaner. Generally, assessments of aquatic invertebrate populations are considered to be among the more reliable gauges of stream health. But some caution is in order in this case.
Prior to the occurrence of large flights several years ago, Hexagenia bilineata was not well known among the species in the mayfly communities of the lower Susquehanna and its tributaries. The native range of the species includes the southeastern United States and the Mississippi River watershed. Along segments of the Mississippi, swarms such as occurred at Columbia-Wrightsville in 2015 are an annual event, sometimes showing up on local weather radar images. These flights have been determined to be heaviest along sections of the river with muddy bottoms—the favored habitat of the burrowing Hexagenia bilineata nymph. This preferred substrate can be found widely in the Susquehanna due to siltation, particularly behind dams, and is the exclusive bottom habitat in Lake Clarke just downstream of the Columbia-Wrightsville bridge.
Native mayflies in the Susquehanna and its tributaries generally favor clean water in cobble-bottomed streams. Hexagenia bilineata, on the other hand, appears to have colonized the river (presumably by air) and has found a niche in segments with accumulated silt, the benthic habitats too impaired to support the native taxa formerly found there. Large flights of burrowing mayflies do indicate that the substrate didn’t become severely polluted or eutrophic during the preceding year. And big flights tell us that the Susquehanna ecosystem is, at least in areas with silt bottoms, favorable for colonization by the Great Brown Drake. But large flights of Hexagenia bilineata mayflies don’t necessarily give us an indication of how well the Susquehanna ecosystem is supporting indigenous mayflies and other species of native aquatic life. Only sustained recoveries by populations of the actual native species can tell us that. So, it’s probably prudent to hold off on the celebrations. We’re a long way from cleaning up this river.
In the absence of man-made lighting, male Great Brown Drakes congregate over waterways lit often by moonlight alone. The males hover in position within a swarm, often downwind of an object in the water. As females begin flight and pass through the swarm, they are pursued by the males in the vicinity. The male response is apparently sight motivated—anything moving through their field of view in a straight line will trigger a pursuit. That’s why they’re so pesky, landing on your face whenever you approach them. Mating takes place as males rendezvous with airborne females. The female then drops to the water surface to deposit eggs and later die—if not eaten by a fish first. Males return to the swarm and may mate again and again. They die by the following afternoon. After hatching, the larvae (nymphs) burrow in the silt where they’ll grow for the coming year. Feathery gills allow them to absorb oxygen from water passing through the U-shaped refuge they’ve excavated.
Several factors increase the likelihood of large swarms of Great Brown Drakes at bridges. Location is, of course, a primary factor. Bridges spanning suitable habitat will, as a minimum, experience incidental occurrences of the flying forms of the mayflies that live in the waters below. Any extraordinarily large emergence will certainly envelop the bridge in mayflies. Lights, both fixed and those on motor vehicles, enhance the appearance of movement on a bridge deck, thus attracting hovering swarms of male Hexagenia bilineata and other species from a greater distance, leading to larger concentrations. Concrete walls along the road atop the bridge lure the males to try to hover in a position of refuge behind them, despite the vehicles that disturb the still air each time they pass. The walls also function as the ultimate visual attraction as headlamp beams and shadows cast by moving vehicles are projected onto them over the length of the bridge. Vast numbers of dead, dying, and maimed mayflies tend to accumulate along these walls for this reason.
The absence of illumination from fixed lighting on the deck of the bridge reduces the density of Great Brown Drake swarms. Some communities take mayfly countermeasures one step further. Along the Mississippi, some bridges are fitted with lights on the underside of the deck to attract the mayflies to the area directly over the water, concentrating the breeding mayflies and fishermen alike. The illumination below the bridge is intended to draw mayflies away from light created by headlamps on motor vehicles passing by on the otherwise dark deck above. Lights beneath the bridge also help prevent large numbers of mayflies from being drawn away from the water toward lights around businesses and homes in neighborhoods along the shoreline—where they can become a nuisance.
Lights out on the Columbia-Wrightsville bridge. Dousing the lights to eliminate fixed illumination on bridges is an effective method of reducing the density of Hexagenia bilineata swarms.With the bridge lights darkened, male Great Brown Drakes, their cellophane-like wings illuminated by headlamps to appear as white spots on the road, number in the hundreds instead of hundreds of thousands in swarms on the bridge near the east and west shorelines.Swarms of Great Brown Drake mayflies are still present at the Columbia-Wrightsville bridge, they’re just not concentrated there in enormous numbers. Evidence includes their bodies found in cobwebs along the entire length of the span.The aptly-named Bridge Orb Weaver (Larinioides sclopetarius) constructs webs along the entire length of the Columbia-Wrightsville bridge, and on many of the buildings at both ends. The abundance of victims tangled in silk must overwhelm their appetite, or maybe they actually consume only the smaller insects. They have their choice. Of the Bridge Orb Weaver, Uncle Ty Dyer says, “When you live along the river, it’s your friendly neighborhood spider, man.”The native Eastern Dobsonfly (Corydalus cornutus) is among the reliable indicators of stream quality in the Susquehanna at the Columbia-Wrightsville bridge. Winged adults, which live for about a week, are clumsy fliers attracted to lights. The aquatic larvae are known as hellgrammites, which require clean flowing water over rocky or pebbly substrate to thrive. Two adults were found on the bridge last evening. It would be encouraging to find more. Maybe we’ll stop back to have another look when the lights are back on.
SOURCES
Edsall, Thomas A. 2001. “Burrowing Mayflies (Hexagenia) as Indicators of Ecosystem Health.” Aquatic Ecosystem Health and Management. 43:283-292.
Fremling, Calvin R. 1960. Biology of a Large Mayfly, Hexagenia bilineata (Say), of the Upper Mississippi River. Research Bulletin 482. Agricultural and Home Economics Experiment Station, Iowa State University. Ames, Iowa.
McCafferty, W. P. 1994. “Distributional and Classificatory Supplement to the Burrowing Mayflies (Ephemeroptera: Ephimeroidea) of the United States.” Entomological News. 105:1-13.
