Alderflies and Dobsonflies (Megaloptera) – LIFE IN THE LOWER SUSQUEHANNA RIVER WATERSHED

May 19, 2026May 25, 2026

The Stewards of Inland Shorebird Habitat

As they travel between coastal wintering ranges and breeding territories in Canada and Alaska, vast numbers of shorebirds pass through the lower Susquehanna region each spring and fall—though few stop here to rest, feed, and provide us with an opportunity to observe them. Prior to the construction of man-made dams and other alterations on our lotic (flowing) waterways including the river, shorebirds took advantage of lateral bars, stream deltas, and other alluvial deposits as places to loaf and re-energize. Before they were drained and filled, some of the valley’s wetlands probably included sparsely vegetated flats where shorebirds could drop in for a brief visit. Previous to their extirpation from our region, North American Beavers were the primary providers of quality habitat for shorebirds and other migrating waders on our lotic waters. Their widespread network of dams, pools, and marshes maximized floodplain function by keeping streams thoroughly connected to their wetlands, nurturing plant communities that not only provided food and shelter for the beavers and other wildlife, but provided superb buffering against erosion while protecting against sediment and nutrient imbalances in lower Susquehanna waterways.

Beaver Dam and Lodge — Beaver dams need not be large, particularly on low-gradient streams where a structure like this is sufficient to create a pool with depth adequate for building and maintaining a lodge and transporting leafy branches and other food items by water.

A beaver lodge assembled in a pool with less than three feet of water, deep enough to provide the family with a measure of protection against terrestrial predators.

Beaver Pool — Beaver pool ecosystems provide homes for hundreds of species of plants and animals, including migrating shorebirds and other waders.

Mud flats in the margins between emergent shrubs and herbaceous plant growth attract migrating shorebirds like this Solitary Sandpiper to the abundance of invertebrate life. Seasonal movements of migrating shorebirds regularly coincide with the reductions of water levels in beaver pools which typically occur between May and September each year.

During its southbound migration in September, a Least Sandpiper searches for arthropods and annelids as it visits a food-rich puddle along the periphery of a beaver pool.

A Pectoral Sandpiper feeds in shallow water near a stand of Common Cattails in a beaver pool.

During migration, rails including the Sora are attracted to dense emergent vegetation in beaver pools. Some will nest in these rodent-managed refugia.

Green Herons visit beaver pools during migration and, due to the reliability of the food supply, will often nest in their vicinity.

So how did this happen? How did the North American Beaver become a keystone species—an animal upon which the majority of other life forms within its ecosystem are so reliant? Well, it’s largely due to the fact that our beavers aren’t particularly fond of a constant stream of noise. More specifically, they don’t like the sound of running water in places where they intend to build and maintain a lodge. And so, as they begin to place sticks, mud, limbs, stones, and other materials within a noisy riffle on a stream, they create a dam, and behind it a pool—a pool that is particularly advantageous for protecting their home and providing a means of conveyance for their construction materials and food supplies.

Cascading Series of Beaver Pools — On a high-gradient segment of stream, beavers will create a cascading series of pools. Because water filters through a beaver dam instead of spilling over it, the work of these meticulous rodents soon silences the sounds of water changing altitude. No more sonorous riffle.

Beaver Dam — **Quiet please.** High seasonal stream flow and damage from storms may create areas where water begins to erode the structure of the dam. Where this condition persists, an adult beaver will soon mend the breach, just to quiet things down. Why would beavers demand such a hush upon their domain? Well, they have poor eyesight, but their hearing is excellent, and they rely upon it to detect danger.

The Louisiana Waterthrush (Parkesia motacilla), one of our earliest-arriving warblers, nests in forests along clear, high-gradient streams. In mid-April, we found this individual and three others squabbling over a breeding territory adjacent to the series of cascading beaver pools shown in the previous images.

Brook Trout and Blacknose Dace — Native denizens of coldwater streams, neither the Brook Trout nor the Eastern Blacknose Dace has any difficulty finding its way through the voids in beaver dams to ascend and descend the sequence of pools.

Beaver Dam and Pool on Low-gradient Stream — On a low-gradient segment of stream, a dam just over a foot in height may be sufficient to create a beaver pool of considerable size. Resembling the water-logged muskeg of the far north, well-established beaver pools form boggy habitats familiar to migrating shorebirds.

Water levels in the pools usually drop along with the stream’s base flow as soon as the effects of the spring thaw and rain showers subside. Feeding areas in shallow water and on muddy ground are often revealed just in time for northbound shorebirds and other waders to stop by in late April and May. These conditions often persist through the growing season and the fall migration of these birds which begins as early as late June and sometimes extends into October.

Tree Swallow at Natural Cavity — Where the pool inundates standing timber such as Red Maple and other species not used by beavers, the dead snags provide vital feeding areas for birds and other wildlife. Cavity nesters like this Tree Swallow seldom find suitable natural housing elsewhere.

Eastern Newts Among Alderfly Exuvia — The mosaic of marshlands and braided stream channels within the beaver pool complex supports an abundance of aquatic life including these breeding Eastern Newts, seen here surrounded by the exuvia left behind following a massive hatch of alderflies (Sialinae). Alderflies are a stream inhabitant during their larval stage and are indicator of clean water conditions.

**A modern-day example of the way fully functional stream floodplains used to look in the Lower Susquehanna River Watershed.** Though the pools may appear pond-like during the cooler and wetter months of the year, by summer the water levels behind the beaver dams recede as the base flow of the stream wanes. Now the true nature of the marsh, shallow pool, and braided stream complex is revealed. Unlike most man-made dams that set a fixed pool elevation regardless of flow by discharging water over the top of the structure or through a spillway or gate, beaver dams merely throttle the flow through their porous construction. Unless the beaver begins plugging small leaks as fast as the stream flow ebbs, the water level in the pools will drop. So they’re not left high and dry, lodges are located in the deepest pools, usually in close proximity to the dam and/or one of the stream courses. Though not inundated during the dry season, the soil in the pool complex is almost always damp and plants grow vigorously there, sequestering nutrients and retaining sediments in beneficial deposition patterns that actually inhibit erosion of the riparian landscape. These streams and floodplains retain their hyporheic zone and freely exchange water with the underlying water table and aquifer. It’s the ultimate floodplain management system, and the beavers don’t even know their doing all the work. (Google Maps base image)

Where another high-gradient segment of stream enters the main pool complex, beavers have created an additional series of cascading pools. The impoundments created by these dams help diffuse stormwater energy and process more nutrients and sediments within the floodplain’s wetland vegetation system. (Google Maps base image)

Sandhill Cranes in Beaver Pool — Beaver pools often refill as stream flows increase following autumn rains. The stockpiles of vegetative foods that grew within the beaver’s domain through the summer then become flooded and are a prime source of nutrition for not only the beavers, but waterfowl during both their autumn and spring migrations. Gleaning and probing Sandhill Cranes often find these habitats to their liking as well.

While North American Beavers have returned to the region, most live as “bank beavers”, residing in the river and larger creeks of the valley where they excavate shelters among the roots of Black Willows and other shoreline trees and shrubs. Floodplain encroachment, legacy sediment deposits, and just plain human intolerance have all conspired to prohibit North American Beavers from performing their magic on smaller local streams. For migrating shorebirds, this continued absence of beaver dam ecosystems has turned much of the lower Susquehanna valley into “flyover country”. Those travelers that do stop to rest and feed concentrate at the few favorable locations such as the lateral bars and the hydroelectric dam-created delta at the Conejohela Flats on the river in Lancaster County. But centralization has its drawbacks. Migrants spending time at concentration points may have a greater chance of contracting and spreading disease. Protracted heavy foraging can degrade these habitats. And over time, features such the lateral bars and delta deposits, including those on the Conejohela Flats, transition into other habitats—riparian forests. A more widespread selection of favorable stopover points for shorebirds, waders, waterfowl, and other migrants is certainly desirable.

IMITATION IS THE SINCEREST FORM OF FLATTERY

Until public sentiment sways in favor of the North American Beaver, wildlife managers are mimicking some of the attributes of their sound-inspired installations.

New Impoundment at Middle Creek Wildlife Management Area — Created by excavating a depression in heavy clay soils, this new impoundment at the Pennsylvania Game Commission’s Middle Creek Wildlife Management Area collects water directly from rainfall and from surface runoff. Its depth at no point is greater than about two or three feet. During the driest times of the year, this space will be a mudflat, and a haven for migrating shorebirds.

Shallow-water conservation impoundments designed, constructed, and managed for migratory waterfowl and waders including shorebirds are not what we typically refer to as ponds—though they are lentic (still) waters. Similar shallow freshwater impoundments at our National Wildlife Refuges are referred to as pools by the United States Fish and Wildlife Service, but smaller versions like this example at Middle Creek very closely resemble the prairie potholes created by glacial scour in the north-central United States and adjacent portions of Canada. Many populations of migratory birds are familiar with pothole ecosystems and, like the beaver pools and marshes, have relied upon them as waypoints along their journeys for centuries.

Pond and Lake Zonation — Impoundments most beneficial to migrating waterfowl and waders including shorebirds are shallow in depth. They lack the deeper waters of a pond or lake and thus have no limnetic (open water) or aphotic zones. Managed throughout as a littoral zone, impoundments grow plants in shallow water or on damp soil through the summer months to chew up nutrients accumulated in waste deposited by visiting birds. This vegetation is then flooded from late fall through early spring as forage habitat for migratory waterfowl. The timing of the fluctuations in water levels approximates those of the beaver pools and marshes on lotic (flowing) waters.

Impoundment Gate — A gate assembly is a water control structure installed to provide seasonal management of the water levels in a conservation impoundment. More than once, beavers have heard the sounds of tumbling waters inside these types of devices and tried to dam them up!

You’ve heard the line, “If you build it, they will come.” Well, it’s true. Here is a sample of the activity witnessed during the past two weeks at the new impoundment completed just several months ago at Middle Creek Wildlife Management Area along the tour route just beyond the “Stop 3” overlook.

The ubiquitous Snow Geese were among the first migratory occupants of the new impoundment. A few are currently lingering in the vicinity.

Ducks soon followed. These Blue-winged Teal were among the last to pass through earlier this month.

Wilson's Snipe — Wilson’s Snipe were among the first species of migratory shorebirds to visit the new impoundment at Middle Creek…

…as were Killdeer, a species which nests nearby.

Then, earlier this month, flocks of shorebirds including these Least Sandpipers were arriving to feed and rest.

Least Sandpiper and a Semipalmated Sandpiper — A Least Sandpiper (left) and a Semipalmated Sandpiper (right).

White-rumped, Least, and Solitary Sandpipers — Here’s a Solitary Sandpiper (right) feeding alongside a Least Sandpiper (center) and a White-rumped Sandpiper (left).

Least Sandpiper and a White-rumped Sandpiper — A Least Sandpiper (left) and a White-rumped Sandpiper (right).

Rare in our area during spring, the White-rumped Sandpiper (Calidris fuscicollis) flies north using the central flyway, then heads south along the Atlantic flyway in the fall, when it tends to be more regular here.

Rocking and teetering along as it looks for food, the Spotted Sandpiper may be one of the easiest of shorebirds to identify.

Chunky little Dunlin (Calidris alpina) with their conspicuously down-curved bills are another easy-to-identify species, particularly in the spring when their breeding (alternate) plumage includes a black belly.

Black-bellied Plovers acquire a handsome set of plumage in the spring as well.

The Semipalmated Plover doesn’t have nearly the flair for ornament its close relatives the Killdeer do; these little shorebirds wear only one ring around their neck instead of two. Think this plover is cute?,…

Speaking of legs, here’s one of dozens of Lesser Yellowlegs that visited the new impoundment during their recent northbound travels.

Though less numerous than the smaller Lesser Yellowlegs, a Greater Yellowlegs seldom goes unnoticed when dropping by the man-made pothole habitat.

Wilson's Phalarope — Everyone likes to see something unusual every now and then, and this impoundment delivers. Just yesterday, we photographed this migrating Wilson’s Phalarope (top center) among two Least Sandpipers (top left and right), a Lesser Yellowlegs (left and slightly forward of the phalarope), and a White-rumped Sandpiper (foreground).

Species of wildlife in addition to shorebirds and waterfowl have already found the new impoundment favorable…

Pairs of breeding Spot-winged Gliders (Pantala hymenaea), seen here in tandem while ovipositing, were swarming the impoundment after arriving earlier this week. These dragonflies are ofttimes unpredictable nomads and are similar in appearance to the usually more numerous Wandering Gliders. To recolonize seasonal portions of their range, both are famous for hitching a ride en masse on storm systems. They share the behavior of finding ephemeral and new bodies of water favorable for egg laying due to their low density of aquatic predators.

We watched this Snapping Turtle arrive to apparently find the newly created waters to its liking. Snapping Turtles are important consumers in a wetland ecosystem. Larger specimens may fill the role of an upper trophic level or apex predator, eliminating vulnerable mid-level consumers including other Snapping Turtles.

Because it bounces its tail up and down like a Spotted Sandpiper, you may at first glance mistake an American Pipit for a shorebird. Long known as the Water Pipit, these songbirds have been visiting the impoundment while migrating north and stopping to feed in nearby croplands.

Seen here with Least Sandpipers is a visiting Glossy Ibis.

Western Cattle Egrets visiting Middle Creek this spring have been frequenting the new impoundment.

Western Cattle Egret — With still little in the way of insects such as grasshoppers available in the surrounding landscape,…

…cattle egrets are looking to find prey like this Green Frog.

Managing saturation levels in shallow-water impoundments to resemble the seasonal variations in beaver pool and marsh systems can create lush growth and wildlife-rich environments. Take a look at some images from a project in a headwaters area of a tributary to Conewago Creek (west)…

Well-established Shallow-water Impoundment — By late July, southbound shorebirds were already using these mudflats to feed and rest. Other sections of the impoundment were dense with emergent and aquatic plants, the latter kept hydrated in deeper pools of the project by the inflow from several captured springs that supplement direct rainfall and sheet runoff to supply its water. During a seasonal drawdown, the exposure of the impoundment’s soils to direct sunlight can provide a measure of disinfection to reduce the chances of disease transmission among its populations of visiting birds and other animals.

Water Lily Growth — In the deeper pools of the impoundment, water lilies and other aquatic plants grow in lush mats to provide cover and feeding areas for resident populations of breeding reptiles and amphibians.

Immature Little Blue Heron — …and this immature Little Blue Heron, a wanderer typical of more southern latitudes.

While walking the road among tall grasses in the supporting landscape surrounding this impoundment, we were at first startled when these Sandhill Cranes strode by going the other direction. We quietly kept moving,…

Sandhill Cranes with Colt — The hatching of this colt is testimony to the vital role wetland ecosystems play in the lives of hundreds of species. Whether they be beaver pools and marshes on lotic waters or man-made shallow lentic waters, each of these habitats is filling a void that left floodplains and other critical lowland biomes faltering. While they can’t replace the full-function floodplain management provided by an active beaver colony, shallow-water impoundments can provide relief for habitat-starved populations of the animals and plants that rely upon them. A constellation of these projects on lands public or private across the lower Susquehanna watershed could help provide refuge for many of our flora and fauna with the most desperately fragmented of ranges.

So that you can relax while observing the comings and goings at a pair of the lower Susquehanna valley’s man-made impoundments, the Pennsylvania Game Commission has erected two viewing pavilions for public use on its lands…

The Haldeman Island observation pavilion is located on State Game Lands 290 just upstream from the Juniata River’s mouth on the Susquehanna at Clark’s Ferry in Dauphin County.

Haldeman Island Pavilion — It overlooks not only the island’s man-made shallow-water impoundments and neighboring grasslands, but the tower used in the 1970s to reintroduce Bald Eagles from Saskatchewan to the lower Susquehanna. Interpretive signs explain the conservation stories of habitats and the eagle reintroduction program.

Middle Creek W.M.A. Observation Pavilion — The observation pavilion at Middle Creek Wildlife Management Area is of similar construction to the one at Haldeman Island. It is accessed from the parking pull-off along the tour road at its intersection with Chapel Road, just before the right turn and incline that leads to the “Stop 3” grassland overlook.

It too includes numerous interpretive signs to help visitors understand impoundment management.

During the next two weeks, the exodus of migrating shorebirds now staged and feeding upon Atlantic Horseshoe Crab eggs on Delaware Bay will commence. During the evening of their departure from the bay, many of these birds cross portions of the Lower Susquehanna River Watershed, particularly east of the river. Stormy weather and other climatic conditions may force some of them to seek a place to put down temporarily, so keeping a close eye on the new pothole-like impoundment at Middle Creek may be a prudent move. After that, waders known as “post-breeding wanderers” can show up at any time. Then, beginning as early as late June, shorebirds begin moving south on a migration that can provide us with viewing opportunities into September and beyond. See you out there!

March 25, 2023April 17, 2023

Underwater View of Life in a Vernal Pool

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.

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…

- - The Pennsylvania Natural Heritage Program page

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…

- - The Pennsylvania Natural Heritage Program’s Vernal Pool Resources page

**Wasted Effort-**A pair of Wood Frogs mating in a dried-up vernal pool.

July 15, 2019July 16, 2019

Friendly Neighborhood Spider, Man

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.