Migrating Waterfowl on Lotic Fresh Waters

Lotic vs. Lentic Freshwater Ecosystems — Some comparative attributes of unspoiled lotic vs. lentic freshwater ecosystems. Low-gradient (slow-moving) lotic waters often create, and remain connected to, accompanying riverine wetlands (a lentic freshwater ecosystem). These swamps, marshes, and ditches absorb, purify, and infiltrate flood waters while supporting a diverse number of plant and animal species.

We frequently perceive all waterfowl migration to be synchronized with the conspicuous movements of familiar species like Snow Geese, Tundra Swans, and Canada Geese—big flights coming south in October and November, then a return to the north in late February and March. And we’re all quite aware of the occurrence of large gatherings of some of these migrants while they make stopovers on some of our largest lentic (still) waters—the man-made lakes and reservoirs created by damming local streams. But did you know that there are populations of colorful waterfowl with dynamic migrations that extend throughout the winter and early spring with movements that are often continuous. Under favorable conditions, these birds favor the lotic (flowing) waters of the river and its larger tributaries as they transit the lower Susquehanna valley. That’s because unpolluted lotic freshwater ecosystems support a greater diversity of plants and animals than lentic waters, and therefore offer more opportunities for hungry migrating waterfowl to find food. Let’s have a look at some of the species that visit the river during their seasonal journeys…

Ice on the Susquehanna — While the urge to head south in the autumn is largely stimulated by the shortening of the photoperiod, it is the presence of ice, particularly on glacial lakes throughout the lands to the north of the lower Susquehanna River basin, that pushes many diving ducks to finally make their way south toward the guarantee of open-water feeding areas along the coast. This movement may occur at anytime between November and late February or, as we have seen during some of the mild winters of recent decades, it may scarcely be noticed at all.

Common Mergansers often lead they way when it comes to migratory diving ducks. They regularly move south in conspicuous numbers by late November and December and are regularly pushing north as soon as the ice begins to melt. During a typical year, it is not unusual for some populations of these large diving birds to remain north of us during the winter. Then, in the days after a sudden rush of frigid polar air, an appreciable increase in ice cover will force a mid-winter movement of birds down the Susquehanna.

Heat Flux Sources Impacting Temperature in Lotic Fresh Waters — Temperatures in lotic fresh waters vary over the length of the stream or river. They are largely determined by the collective impact of the numerous sources of heat flux depicted in this graphic. (Environmental Protection Agency image)

Buffleheads begin passing through the lower Susquehanna region in Novemeber on their way to coastal saltwater bays for the winter. Lingering populations feed by diving in the river’s pools and riffles for benthic invertebrates including snails and insect larvae. Lesser quantities of aquatic plant matter supplement their diet.

Many Common Goldeneyes will remain on shallow, ice-free waters of the northern lakes and rivers sculpted by the most recent glacial event, but only until they are forced south into and through the lower Susquehanna valley by the encroachment of freezing conditions. On the river, they are among the dozen or so species of diving ducks we see visiting or passing through during the typical late fall and early river.

Geomorphological Zonation of a Stream or River — During their visits to the lotic (also known as riverine) fresh waters of the Susquehanna and its largest tributaries, benthic-feeding waterfowl make short dives to take advantage of the plants, small fish, invertebrates, and other food sources inhabiting the stream bottom in the riffles and pools of the free-flowing waterway. Substrates, listed here by size (in descending order), along with other parameters influenced by this zonation determine the variety and abundance of the forage available to migrating waterfowl and other consumers. Ice or high water and poor visibility due to flooding can render the riffles and pools of the channel unusable for feeding. The birds then must choose to either linger and rest without feeding or leave the lotic freshwater habitat to seek sustenance. During a flood, this may require relocation to a nearby lentic (still) body of fresh water such as a lake or reservoir. The presence of ice will almost invariably force the birds to fly on to the Atlantic Coastal Plain and the tidal waters of its bays and estuaries.

Hooded Mergansers are one of the few species of diving ducks likely to utilize flooded shoreline timber and riverine (fluvial) wetlands as refuge from high water on Susquehanna.

Pied-billed Grebe and Canvasbacks — A Pied-billed Grebe and a pair of Canvasbacks on an ice-free stretch of the lower Susquehanna in mid-winter feed and loaf in a riffle-flanked pool where a large mat of American Eelgrass, a submerged aquatic plant also known as Tapegrass or Wild Celery, grows during the summer. The complex layout of riffles and pools in a river this size offers tremendous opportunities for a diverse array a species to find their niche wherein they can survive and flourish.

The presence of ice forces Buffleheads and other diving ducks to gather in turbulent open water, often below a riffle or dam. Another alternative is to continue on toward the salty bays and estuaries of the coast. High water may push these birds into the shallows among the flooded woods to feed, but they seldom utilize heavily forested riparian wetlands as a refuge due to their need for a running start to get airborne.

**Riffle and Pool Characteristics of High Gradient (A) and Low Gradient (B) Streams**. This graphic illustrates the change in deposition characteristics of a stream or river as its gradient decreases. A high-gradient stream (A) has a rapid velocity, often forms falls, and tends to carry away a high volume of all but the largest of particles of potential substrates as they erode from the surrounding landscape. On a low-gradient stream, the loss in water velocity reduces the water column’s ability to transport even the smallest of substrate particles. Deposition of this gravel, sand, silt, and clay forms lateral bars that over time create the familiar meandering path of a naturally flowing lowland stream. (National Park Service image)

Benthic Life in Lotic Freshwaters — Benthic substrates in lotic freshwater pools and riffles support an abundance of life forms ranging from colorful diatoms on rocks and cobble, to invertebrates including snails and insect larvae, to fishes like this young Channel Catfish. Free of accumulations of sediment, this river bottom not only provides habitat for a healthy fishery, it facilitates the bidirectional exchange of water between the Susquehanna and its underlying aquifer.

On the lower Susquehanna, populations of young Quillback suckers are found almost exclusively in clear, high-gradient pools.

The Harlequin Duck winters along the rocky shores and man-made jetties of the Atlantic coast. In summer, they nest on fast-moving, headwater streams well to our north. Very rare on the Susquehanna, this is the first of two individuals found during March and April of 2025. It was observed feeding in the swift waters of the high-gradient riffles and pools where the river cuts through Blue Mountain north of Harrisburg. During previous weeks, Harlequin Ducks were being seen along the coast as far south as the mouth of the Chesapeake at Cape Charles, Virginia. It’s very possible that some of these birds traveled north through the bay area and up the Susquehanna on their way north.

**Fluvial Geomorphology of a Stream**. Many of the Susquehanna’s tributaries pass through each of these three erosional zones. Along the way, they carry out the process of breaking down the mountains formed by the Allegheny orogeny, the collision of North America and Africa that created the supercontinent Pangea about 325 to 260 million years ago (during portions of both the Carboniferous and Permian periods). Today’s main stem of the lower Susquehanna passes through a transfer zone (Zone 2), carrying eroded materials to a depositional zone (Zone 3) located within the ancient Susquehanna canyon stretching from Havre de Grace, Maryland, to Norfolk Canyon on the edge of the continental shelf. Within this zone, more than a 10,000-year accumulation of post-glacial sediments lies submerged by the rising waters of the Atlantic Ocean and Chesapeake Bay. Although the present-day lower Susquehanna is largely a transfer zone, some deposition occurs along low-gradient segments of the river, particularly where its course parallels the watershed’s ridges both above and below the high-gradient rapids where its path has eroded passage through the highlands. (National Park Service image)

Although the present-day lower Susquehanna is largely a transfer zone, some deposition occurs along low-gradient segments of the river, particularly where its course parallels the watershed’s ridges both above and below the high-gradient rapids where its path continues to erode passage through the bedrock. On this 1908 map of the Susquehanna at Conewago Falls, alluvial terraces of gravel, sand, silt, and clay can readily been seen as pale areas nearly lacking brown contour lines along the shorelines and islands of the river. Deposits within most of these terraces date back to the melt period following the most recent glacial event and beyond. The delta shown at the mouth of Conewago Creek (west) includes massive volumes of material deposited by both the creek and the river. This delta is currently known as Brunner Island, much of it developed as the site of a coal/gas-fired electric generating station. Terrace deposits along the Susquehanna’s shorelines created extensive perched marshes and swamps (wooded marshes) fed by rains, high river water, small streams, and springs, the latter often seeping from the base of the rocky escarpments carved by the ancient Susquehanna and defining the present-day inland border of the floodplain. We call these sites “Alluvial Terrace Wetlands”. Few of these critical components of river morphology survive. Those not drained for farmland were obliterated by urbanization and canal, railroad, and highway construction. (United States Geological Survey base image: Middletown, PA, quadrangle, 1908)

Water Willow is a familiar emergent plant that colonizes lateral bars and other alluvial deposits in low-gradient segments of the Susquehanna.

Water Willow Roots — The fine roots of Water Willow collect sediment and absorb nutrients while creating dense cover for young fish and numerous species of invertebrates including the Virginian River Horn Snails seen here.

Prior to the nineteenth century, the low-gradient flow regime of the river both above and below the riffles at Chiques Rock (lower right on map) created prime wildlife habitat. The natural accumulations of nutrients and substrates carried into and through the lotic waterway’s pools and riffles were cycled into an ideal growing medium for extensive mats of American Eelgrass and other aquatic plants. This underwater forest hosted a seemingly endless abundance of invertebrates and fishes (both resident and migratory)—supporting a variety of consumer species including various populations of humans. But soon after the mass clearing of much of the watershed’s land for farming and lumber, the mill ponds created by dams constructed on streams to power saw and grain mills became brimful with sediments eroded from the unprotected ground. During storm events, torrents of these sediments then flowed full bore toward the Susquehanna, and began accumulating in the low-gradient segments of the river.

Legacy Sediments — Sediments left behind after the removal of mill dams are known as legacy sediments. They disconnect the stream from its historic floodplain and riverine wetlands, thus intensifying the impact of high water in the surrounding landscape. As these nutrient-charged deposits wash away, they become a source of pollution in the waters of the Susquehanna and Chesapeake.

A mat of American Eelgrass growing in the flowing waters of the Susquehanna below Conewago Falls. Eelgrass and other submerged aquatic plants consume nutrients deposited in the cobble, gravel, and sand substrate of the river’s pools. Excess quantities of smaller particles of silt and clay can clog the substrate and thus inhibit the hyporheic exchange of water between the stream channel and the underlying aquifer, often diminishing the biomass and diversity of organisms inhabiting this benthic habitat. Buried in these life-choking sediments, the river bottom becomes inhospitable to growth of submerged aquatics including eelgrass.

This low-gradient stretch of the Susquehanna at Marietta flows parallel to the Chickies Quartzite “Hellam Hills” before making a sharp right turn to punch through the ridge as a series of rapids. Formerly a fully functional lotic ecosystem and a paradise for migrating waterfowl, this river segment is now impaired by accumulations of nutrient-laden sediments from agricultural and urban runoff.

Conewago Falls Flood Waters — Nutrient and sediment-loaded flood waters roar over the York Haven Dam and Conewago Falls…

Flood Waters at Haldeman Riffles — …past Brunner Island (left power plant stacks) and through Haldeman Riffles and the Shocks Railroad Bridge…

Flooding on the Marietta Broads — …into the stretch of river along the base of the “Hellam Hills” known as the “Marietta broads”. The low stream gradient here produces a slower current and increased deposition of sediments.

Redheads, Canvasbacks, and a Horned Grebe — Anytime from November to April on the lower Susquehanna, a group of Redheads, Canvasbacks (3 birds to right of the middle of the picture), and a Horned Grebe (lower left) is a welcome sight in a riverine pool known to have a summertime growth of American Eelgrass. Noted Dr. Herbert Beck in 1924 when describing the Canvasback, “Like all ducks, …, it stops to feed within the county (Lancaster) less frequently than formerly, principally because the vast beds of wild celery which existed earlier on broads of the Susquehanna, as at Marietta and Washington Borough, have now been almost entirely wiped out by sedimentation of culm. Prior to 1875 the four or five square miles of quiet water off Marietta were often as abundantly spread with wild fowl as the Susquehanna Flats are now. Sometimes there were as many as 500,000 ducks of various kinds on the Marietta broad at one time.”

Aythya genus ducks. — Today, seeing just dozens of Aythya genus ducks (Redheads, Canvasbacks, scaup) on the lower Susquehanna is a notable event. If they happen to be forced down by inclement weather while migrating through, you might get lucky enough to see several hundred.

During the past two centuries, as food supplies in the Susquehanna grew increasingly compromised for benthic feeders like these Lesser Scaup and other diving ducks, a change in distribution was necessary for survival.

Lesser Scaup in flight. — As individual species, Lesser Scaup and other waterfowl that fail to adapt to natural or man-made changes in their habitats and food supplies may see their overall global numbers falter.

Despite being located in the transfer zone, the lower Susquehanna has become a significant depositional zone along much of its length, mostly courtesy of the placement of sediment-trapping man-made dams.

Following construction of the mill dams and ponds on nearly every mile of the lower Susquehanna’s low-gradient tributary streams, enterprising parties moved on to the river. The first significant spans were constructed using wide timber cribs filled with large rock. They were placed to create water deep enough to allow canal boats to cross the Susquehanna at both Clark’s Ferry at the mouth of the Juniata River in Dauphin County and at Columbia/Wrightsville. These dams also diverted water into the newly excavated canals: the Pennsylvania Eastern Division Canal (completed in 1833) which followed the river’s east shore from Clark’s Ferry to Columbia, and the Susquehanna and Tidewater Canal (completed in 1840) along the west shore from Wrightsville to Havre de Grace, Maryland. Placement of these sediment-trapping man-made dams began a process of converting vast mileage of the lower Susquehanna from a transfer zone into a deposition zone. In addition, layout of the canals and locks followed the contours along the base of the riverside ridges, seriously altering most of the alluvial terrace wetlands that the river had created as a feature of its floodplain during the post-glacial period.

Construction of the canal dams was just the beginning. During the twentieth century, more massive dams would be added to the main stem of the river for hydroelectric energy production at York Haven, Safe Harbor, Holtwood, and Conowingo. Upon their completion, the days of unassisted anadromous fish migrations were over. On the river and its tributaries, smaller low-head dams maintain water levels for boating and recreation, but still function as current diminishing pseudo-lentic waters that blanket the lotic riffle and pool substrates with polluted sediments.

MAN-MADE DAMS TURN LOTIC WATERS INTO UNFLUSHED TOILETS