scat – Linda J. Spielman

The Secret Lives of Voles

Unless you own a cat with an inclination toward hunting, you’ve probably never seen a vole. These little rodents prefer to lead their lives out of sight, spending their time hidden in thick grass, tunneling in forest leaf litter and humus, or sheltering beneath the snowpack. They are sought out by many predators, but their secretive habits and prolific rates of reproduction allow them to thrive.

There are a number of vole species, and they all have chunky bodies, short legs, and stubby ears, like the meadow vole pictured in the photo above. As the name suggests, grassy fields and meadows are the domain of meadow voles. They’re also known as meadow mice or field mice, although wild mice (white-footed and deer mice) differ from voles in their lifestyles and dietary habits. Another common vole species is the woodland vole (also called the pine vole), which prefers forests but also frequents orchards. Voles are not to be confused with moles, which are insectivores, not rodents.

When conditions are right, voles do reveal their presence to us. In winter this can happen when the snow forms into icy crusts which the small creatures can’t penetrate. In the next photo the hard crust was covered by a shallow layer of new snow, and voles (probably woodland), could only bulldoze their way through the softer top layer. Why the meandering pattern of grooves? The animals may have been searching for food, or for cavities leading to underground spaces. Woodland voles are more fossorial than meadow voles and often dig tunnels in the upper layers of forest soils.

When winter snows accumulate, voles find safety and plenty to eat under the snowpack. Meadow voles range throughout snow covered fields, creating runs that they use repeatedly as they search for food. The evidence of their travels shows up after the snow melts as grooves in the flattened grass thatch like those in the next photo.

The diet of meadow voles is diverse but usually includes grasses. Cut sections of grass like the ones in the next photo are often found after the snow disappears.

Another kind of evidence to be found in newly exposed vole tunnels and feeding areas is vole scat, which consists of tubular pellets like those in the central part of the next photo.

The construction of subnivean runs involves the removal of dirt and plant material from active runways, and once winter is past these appear as windrows of debris. The ones in the photo below were made by a meadow vole. Woodland voles leave similar piles of dirt and humus on the forest floor.

When the snow is deep voles are able to access an additional food source, the inner bark of woody plants. Large chews made by voles can be seen on the white pine in the next photo–this would have happened when the animals were safely hidden by deep snow. The thin layer of living cambium cells is the only nutritious part, and fragments of the inedible outer bark sometimes accumulate below the chews.

Tiny tooth marks, like the ones on the buckthorn stem in the next photo, can sometimes be seen.

In the warmer months vole sign is harder to find, but if you wander slowly through grassy fields and meadows you may make some interesting discoveries, especially when vole populations are high. Small openings like the one in the photo below are the first clue to the presence of the shy creatures.

Vole runs are usually completely hidden by the covering of grass and thatch, but if you spread the grasses backward from an opening, you’ll expose the corridors used by the voles.

If you’re very alert as you walk through a field, you may find a meadow vole nest. I found the one shown below in mid-May, so it was only partially hidden by new grass growth. Later in the season they’re much harder to find.

In the next photo you see another unoccupied nest that I opened to show the finely divided fibrous lining. Winter nests like this one are well insulated against the cold. Exits to the tunnel network can be seen at the upper left and the lower right.

Vole populations go through cycles of abundance and scarcity, and when numbers are high, changes in behavior may bring the animals into the open. During a population boom you may–if you stand very quietly–catch a glimpse of a vole as it pops out of a tunnel. But whether or not you ever have that experience, there’s always evidence of the presence of these animals to be found if you know what to look for.

Perfect Perches

Perches–they’re important to wild creatures for many different reasons. The gray squirrel in the opening image (from yardandgarage.com) is using a perch as a feeding site. The next photo shows a Norway spruce whose dead lower branches provided feeding perches for a red squirrel. You can see how the piles of inedible cone cores and scales accumulated under the branches the squirrel perched on. These accumulations are called middens, and they can build up over time into substantial mounds.

Favorite perches often show signs of usage. The red squirrel that used the perch shown below left a cone scale and a number of opened seeds, some with wings still attached.

Red squirrels may mark perches by biting them. In the next photo you see a Norway spruce branch that bears the distinctive paired incisor marks made by a red squirrel. The lower branches of conifers are usually dead, so these marks don’t heal over and may last quite a while.

In the next photo you see a discovery I made during summer a few years ago. The Norway spruce cone crop that year was early and abundant, and a red squirrel had left a cone core, stripped of its supply of edible seeds, resting on the perch it had used. And in case you’re wondering, no, I didn’t put it there, it was all the squirrel’s doing. The scales that dropped as the squirrel fed can be seen on the ground at the base of the tree.

The photo below shows an unusually well elevated feeding perch used by a gray squirrel.

In the next photo you see what I found on top of the log: the remains of an acorn the squirrel had fed on.

Here’s a perch used not for feeding but for food storage. A gray squirrel lodged a black walnut in the crotch of a honeysuckle branch. I’m not sure what the squirrel’s motivation was–perhaps it was to keep the walnut away from other squirrels.

A perch doesn’t need to be overly high to be suitable. In the next photo you see a log used by a squirrel–it could have been a red or a gray–feeding on a white pine cone.

Rocks can also make good perches. Last August a red squirrel harvested young larch cones and brought them to the rock shown below for consumption. Where rocks or logs are available they are preferred over ground level feeding sites.

But food isn’t the only thing drawing animals to perches. A red fox balanced on the log in the next photo in order to deposit its scat. Scat is important in intraspecies communication, and wild canines prefer to leave their scat in conspicuous positions. Sometimes this requires a little acrobatic ability to position the scat just right.

One of my favorite spring experiences is hearing the drumming of ruffed grouse. Males in search of mates perch on logs or other raised features and beat their wings to produce a resonant booming sound. They prefer platforms that are unobstructed and raised well off the ground. You can see a spot in the center of the image where the bark was dislodged by the drumming bird.

Perches can also be used as observation posts. In the next photo you see a mound of earth thrown up by a falling tree. There were tracks–they were barely visible so I didn’t include a photo–going up the side. The size of the impressions suggested a fox.

On top of the mound (shown in the next photo) there were obvious signs of disturbance, showing that it had been used as a perch. The fox would have sat quietly while it listened, looked, and sniffed for signs of prey animals.

We seem to have circled back around to the topic of food, so here’s my last example of a feeding perch. A black bear climbed the beech tree in the photo below and pulled a nut-bearing branch inwards until it broke off. The bear consumed the goodies, pushed the branch aside and pulled another one inward until it broke. The discarded branches formed a tangled cluster, and the bear might even have stood on the growing mass of harvested branches as it continued to pull more branches in. These branch clusters are known by the somewhat misleading term bear nests, although they have more in common with squirrel middens than with nests. With healthy beeches becoming less abundant, bear nests in beech trees are harder to find than they used to be, but the same kind of sign occurs in apple, black cherry, serviceberry, and oak trees.

Wild creatures know their territories in minute detail, and they’re familiar with all the best perches. The attributes of a perfect perch vary somewhat with the specific animal and situation, but safety and accessibility are always important. The location also needs to be appropriate to the animal’s purpose, whether it’s to consume food, to find food, or to advertise its presence. If we stay alert for perches we can begin to understand what makes a good perch and what they can tell us about the lives of the animals.

What Goes In Comes Out

A big part of understanding animal lives is knowing what they eat. There’s lots of general information available in books and other publications, but to understand the dietary habits of the animals in one’s own landscape requires a few steps beyond that. We can observe animals when they’re hunting or feeding, and we can interpret chews, feeding sites, and feeding leftovers. But for many mammals, especially omnivores and carnivores, scat is the best tool. Scat contains the undigestible parts of everything an animal ingests, and it remains long after the creature has left the scene.

For herbivores, plant fibers make up the bulk of eliminations, so their scat has a grainy texture like the rabbit scat shown below. When herbivores eat foods high in water content their scat may be darker and softer, but the fibrous essence can still be seen.

Raccoons are omnivores, and their latrines often contain scats with a variety of contents. In the next photo you see raccoon scats with grape seeds and skins, apple seeds, ant parts, deer hairs, and the amorphous remains of deer flesh. An important safety note: raccoon scat may contain the eggs of a parasite that can infect humans, so it should never be touched with bare hands. To be on the safe side, it’s best to use sticks or other tools to manipulate scat, no matter whose it is.

The photo below shows river otter scat filled with crayfish shell fragments. In locations where fish are the main prey item, fish scales and bones will be the most common contents. In coastal marshes scats with crab shell fragments may predominate, indicating that crabs make up the bulk of the animals’ diet.

For many omnivores and carnivores scat contents vary with the changing seasons. The bear scat shown below was photographed in early May, and it’s made up of the remains of the newly emerging leaves and shoots the bear had been eating. Bears lose weight during hibernation and for many weeks afterwards because the grasses, sedges, and young shoots they must subsist on are energy-poor foods.

It’s only when higher quality edibles become abundant that bears begin to put on the pounds. The bear that left the scat shown in the next illustration had been feasting on black cherries. The summer diet of fruits and berries is often supplemented with insects, and you may find bear scats containing ant or yellowjacket parts.

Scats like that pictured below, full of fragments of acorns and hickory nuts, begin to show up in late summer. The seasonal abundance of acorns, nuts, and fruits, as well as increasing insect populations, provides a crucial, energy dense diet. At this time bears transition into a period of hyperphagia, and spend most of their waking hours seeking food or eating. The fat stores they put on will carry them through their winter hibernation.

The scat of canids reveals that their diets also follow seasonal cycles. Winter and early spring fare is mostly made up of animal prey and carrion. Signs of feeding on deer carcasses start to show up during the fall hunting season and continue through the winter. The coyote scat in the photo below contains deer hair and leg bone fragments. Foxes also feed on deer carcasses, but they aren’t powerful enough to crack large bones to get at the marrow the way coyotes do. Deer killed by hunters (and the carcasses resulting from the vehicle collisions that seem to spike during hunting season) may be preserved well enough in the cold to last through most of the winter. Carcasses of winter-killed deer also provide scavenging opportunities.

The red fox scat shown below (photographed in mid-March) contains the remains of a small rodent that was swallowed whole. There’s a leg bone in the chunk at the lower left, a molar in the piece at the top, and an incisor in the segment at the lower right. The bones are embedded in twisted masses of short hairs. Positioning its scat on the manhole cover was the fox’s way of signaling its presence to other foxes in the area. Small rodents and other small mammals are a winter mainstay for foxes and coyotes.

Like bears, canines graduate to summer foods as they become available. A sure sign that berries are in season are finds like the coyote scat shown below, filled with raspberry seeds. Note that the segments are tubular and blunt-ended rather than tapered like scats made up of animal remains.

As summer progresses, the menu widens. The red fox scat in the next photo (found in early September) contains acorn shells, apple skins, and fragments of field corn kernels.

Some scats lead to surprising discoveries. The next photo shows some gray fox scat containing the remains of a frog. Hollow leg bones are clearly visible, and when I pulled it apart I saw the still articulated bones of a rear foot. It’s a bit unusual to find frog remains in fox scat, but the really surprising thing is that I found this in early December. The weather had been mild, and apparently some frogs had not yet gone underground for the winter.

Food is central to survival, and scat can provide direct information about what animals eat and when they eat it. The many stories scat has to tell can illuminate not just feeding habits, but also interactions among animals, and interactions with the surrounding landscape. Each story adds to our connection with the animals around us.

Muskrats: Life in Two Worlds

Water and land: they pose very different challenges to the creatures that inhabit them. And yet some animals manage to live in both worlds. The muskrat is a semi-aquatic mammal, at home in the water and comfortable (although not as nimble) on land. The dome-shaped lodges made by muskrats (seen in the photo below) resemble beaver houses, but are smaller and are made with non-woody plants instead of the woody material used by beavers. When conditions are suitable muskrats make bank burrows with underwater entrances. Unlike beavers, they don’t build dams, and they prefer quiet or slowly moving water. Aquatic and semi-aquatic plants make up the largest part of a muskrat’s diet, but the animals also spend time on land harvesting non-woody terrestrial plants. They are also known to consume aquatic animals, including clams, mussels, crayfish, frogs, and fish.

A common sign of a resident muskrat is scat, usually found in small collections on rocks and logs that protrude above the water. These deposits announce a animal’s territorial claim to the pond, marsh, or stream where they’re found. The latrine shown in the next photo is on a large rock at the edge of a river, and it’s unusually large. The quantity and the combination of fresh and weathered scat indicate that a muskrat was actively patrolling its stretch of river.

On land muskrats generally move at a walk. In the next image the direction of travel is from left to right, and because of the snow the animal to placed its hind feet directly in the holes made by the front feet on the same side. A tail mark undulates between the tracks.

When the footing is more favorable muskrats use an overstep or indirect register walk. The trail in the next photo goes from upper right to lower left. Pairs of prints form an overall zig-zag pattern, and in each pair the rear track lies ahead of the front. The sequence, starting from the upper right, is right front, right rear, left front, left rear, right front, right rear. If you’ve noticed that the front prints seem smaller than the rear prints, you’re absolutely correct.

The difference in size is easy to see in the next photo. The right front track is on the left, and the right rear is on the right (direction of travel left to right). You can see all five toes in the front print–the tiny innermost toe is a little nub on the upper edge of the print just behind the full-size toe ahead of it. The four large toes of the front track are connected to the middle pad, and behind that there are two bumps that make up the heel pad. If these characteristics remind you of small rodent tracks you’re right on target. Muskrats are indeed rodents, although they have diverged from other rodents in many of their adaptations. In the rear track five toes can be seen, although the innermost toe impression is just the tip (above the other four rear toes and to the right of the third toe on the front print). The middle pad of the rear print made a partial impression at the bases of the toes, and–as is often the case–the heel pad did not touch down at all.

If danger threatens while a muskrat is on land, it hurries toward the safety of the water at a modified bound or lope. In the photo below you see a typical muskrat bounding pattern, with the smaller front tracks ahead of the larger and more widely set rear prints (direction of travel toward the top). The muskrat’s front feet slid forward into the soft mud, so the tips of the toes lie hidden in the muck. The larger rear feet didn’t sink as far and all five toes show clearly. Except for the relative positions of the front and rear tracks, this bounding arrangement is, again, reminiscent of the bounding patterns of many small rodents.

Muskrats possess a feature that is–as far as I know–unique among semi-aquatic animals: the toes of the hind feet are equipped with fringes of stiff hairs. In the next photo you see a left rear track, oriented toward the top. (There’s also part of a left front print to the lower right of the rear print.) The toes of the rear track are slender and finger-like, and the hair fringes make shelf-like impressions around them. These hair fringes add surface area and enhance the muskrat’s swimming ability. The smaller and un-fringed front feet are more suited to grasping and handling objects.

As you explore wetlands you’re likely to see swimming mammals, and you may find it difficult to know which creature you’re observing. There are clues that can help, starting with size. The smallest are water shrews and star-nosed moles. I’ve never seen shrews or moles swimming, so I’ll just point out that their size means they probably won’t be confused for anything but each other. Of the larger mammals, the ones whose tracks and sign we’re likely to find, the smallest is the mink. Minks swim with their entire body visible above the water, from head to furred tail. Their bodies are long and slim, their ears protrude from their heads, and their tails can usually be seen gently swaying from side to side on the surface. Muskrats are heavier than minks but their chunky bodies are about the same length. They swim with their heads and bodies showing above the water. The muskrat’s hairless tail is flattened vertically and can be seen undulating from side-to-side at the surface. The ears are small and don’t protrude from the head.

Next in the size progression (going by weight) is the river otter, with a body length of two to three feet and a powerful, furred tail that tapers from a muscular base to a small tip. Otters often swim with just their heads showing above water, but they may also undulate up and down or make short, playful swerves and dives. Their ears protrude noticeably from their heads. Our largest semi-aquatic mammal is the beaver, with a body length about the same as an otter but weighing up to twice as much. Beavers swim with most of the body and the tail below the water surface, and their ears protrude from their heads.

Muskrats are one of our most common semi-aquatic mammals. You may be fortunate enough to observe one in its watery habitat, or you may instead find evidence of the its presence. Either way, take time to contemplate the muskrat’s place in the panorama of living creatures and the adaptations that make it so successful.

Where Do The Bones Go?

Have you ever wondered what happens to all the bones? Animals are dying all the time, and when they die their soft tissues are eaten by predators and scavengers, picked off by birds, ingested by insects, and decomposed by microorganisms. This leaves just bones, like those of a rabbit shown below. But we don’t see bones littering the landscape, so what happens to them?

First let’s consider small animals. When a tiny creature such as a vole is killed by a predator, the catch is swallowed whole and the bones are crushed and partly assimilated. Undigested bone fragments are eliminated in scat (or pellets if the hunter was a hawk or owl). You can see small bone fragments in the red fox scat shown below–there’s also plant material, tiny hairs, and what appears to be a whisker. Scat like this will eventually be weathered and dispersed into the soil. Even if a small animal isn’t completely consumed immediately, its remains will be broken down, dispersed, and probably hidden from our view by its surroundings.

But what of larger animals whose carcasses would be more obvious? Deer immediately come to mind, but the question also applies to bears, coyotes, woodchucks, raccoons, and other similar sized animals. We do occasionally see the remains of recently deceased animals, like the deer carcass in the next photo, but why don’t we see piles of old bones lying around everywhere?

The answer has to do with the nutritional value of bones. The deer femur in the next photo was cracked open by a coyote to get at the marrow. (I say coyote because the only other animal in our region which is powerful enough to break a deer leg bone would be a bear, and there were no bears in the area where the bone was found.) Toward the upper end of the larger piece you can see some striations which were probably made by the coyote’s molars as it worked at the bone.

We sometimes see evidence of the utilization of bones this way in scat. The coyote scat in the next image contains an abundance of deer bone fragments and deer hair. The hair would have cushioned the sharp bone edges and prevented injury to the animal’s digestive system. It wouldn’t take long for bone fragments like these to be hidden in the upper layers of soil.

In addition to marrow, bones contain calcium, phosphorus, and other minerals which may be lacking in the diets of wild animals. Mineral deficiencies are especially likely for herbivores. Many animals supplement their nutrient intake by chewing on bones, and they usually choose less daunting ones such as scapulas, ribs, and vertebrae. The bones of birds, reptiles, and smaller mammals such as rabbits can also be utilized by less powerful animals. Even deer have been observed chewing on bones. This kind of chewing may not leave obvious signs–just ragged edges, missing ends, or random gouges.

Rodents also gnaw on bones, and the evidence of their activity is often more conspicuous. In the next photo you see a segment of deer leg bone lodged on a midden at the base of a Norway spruce tree. Middens, piles of discarded cone cores and scales, are created when a red squirrel repeatedly uses a favorite perch to feed on cones. The red squirrel that claimed this tree must have used the same perch to work on the bone.

In the next photo you can see the grooves made by a squirrel’s incisors as it chiseled off bone shavings.

Smaller rodents, like voles and white-footed mice, leave finer grooves like the ones in the next photo.

These creatures weren’t after marrow, since the bones were relatively old and the marrow had been removed long ago. This behavior is probably driven in part by the need to supplement their mineral intake, but rodents also chew on bones (and antlers as well) to maintain their teeth in good condition. Their incisors grow constantly, and are subject to malocclusion if not shaped and worn down with regular gnawing. The same is true for rabbits and hares, which are also known to gnaw on bones.

As time passes carcasses are pulled apart and bones are cleaned of soft tissue, scattered, broken, crushed, pulverized, chewed, and ingested by many different animals. Rather than piling up as useless cast-offs, animal bones gradually disappear as they are utilized by living creatures. Animals are part of the web of life both while they are alive and after they are dead.

Conspicuous Communication

If you’ve ever found a pile of feces perched in a conspicuous spot, you’ve encountered a message from an animal. Canines are especially likely to communicate this way, and they’ll use any location that makes a good exhibit. The photo below shows red fox scat displayed on the base of a fallen log. There’s both recent and older scat–recognizable by its lighter color–indicating that this location has been used more than once. One older chunk is nestled in the center of the new deposit and another rests below it on a shelf of wood.

Our olfactory abilities are too limited to appreciate the complex bouquet of chemicals in scat, but for canines–and probably other species–each deposit conveys information. The specific content of the communication could be establishment of a territorial boundary, advertisement of mating availability, or reinforcement of group cohesion. Scat can also indicate the health, status, and identity of the animal which produced it. The coyote scat in the next photo was in the center of a road rather than on a raised object, but it’s placement made it noticeable nevertheless. I found this in June, when we would expect coyote parents to be leading their offspring on short explorations, and my best guess is that the message was territorial in nature.

Important locations may accumulate a number of deposits. The rock in the next image must have been significant, because there are four different scats on the rock and several more which fell off to one side and aren’t visible in the photo. All of the deposits were left by red or gray foxes, and the contents include apple skins and seeds, hair and small bones, and insect parts. The most intriguing one is the chunk at the lower right.

A closer look shows that it contains porcupine quills.

An ant mound formed the pedestal for the red fox scat in the next shot. I found it in early spring, so the ants would still have been deep underground when the animal stood on the mound and dropped its feces directly on top.

Manhole covers can provide suitable display locations. The red fox that left the scat in the next photo had dined on a small rodent, as indicated by the short hairs and small bones it contained. The manhole cover was in a grassy trail and allowed the scat to stand out in the uniformly green surroundings.

Sometimes scat seems to represent an assertion of confidence. Coyotes will kill foxes, so the smaller canines are usually careful to avoid encounters. In the photo below a recent gray fox scat (at the lower right) sits on an older accumulation of coyote scat. The deer hair in the coyote scat shows that the animal had scavenged on a mostly cleaned out carcass, while the gray fox had eaten meat from a fresher carcass.

Any protruding object is a potential platform for canine scat. The photo below shows a deposit of coyote scat on a pile of horse dung.

In the next photo you see one of my most surprising finds. A gray fox had deposited scat on top of a rock cairn which marked a trail junction. This must have required a delicate balancing act, because the pile of rocks was tall enough that the fox would have needed to place at least one rear foot on the cairn.

The conspicuous locations often chosen by wild canines mean that we often notice the scat left by wild canines. We’re less adept at interpreting the messages contained therein. But even if we miss what’s most important to the animals, it’s fun to enjoy the creative and sometimes whimsical positioning of the scat of foxes and coyotes.

Streamside Discoveries

As the high water levels of late winter and early spring subside, stream and lake margins become interesting tracking locations. Water is a magnet for wildlife, and visiting creatures leave the evidence of their activities along the shoreline. A great blue heron left the collection of tracks shown in the photo below. The feet of herons resemble the feet of songbirds, with one backward-pointing toe and three forward-pointing toes. But unlike most songbirds, the toes of herons don’t all meet at one point. There’s a left print (facing toward the lower right) in the upper left corner of the photo that shows this nicely. The junction between the backward-pointing toe and the inner forward-pointing toe lies to the left of the intersection between the two outer toes. Another way of saying this is that the two outer forward-pointing toes join a little to the outside of the center of the foot. The same asymmetry shows in the right track in the lower right corner.

The spotted sandpiper is another bird that patrols stream and lake margins. These small birds–about the size of a starling–search for invertebrates on the edges of streams, ponds, marshes, and other bodies of fresh water. Their tracks (shown in the next photo) reflect their erratic and meandering movements. The three forward-pointing toes are relatively symmetrical and diverge at wide angles. On the back of the foot there’s short spur oriented to the inside that may or may not make an impression in tracks. The left print just below the stick in the upper right corner shows the spur nicely.

Raccoons prefer comfortable surfaces so it’s no accident that the animal that left the tracks shown in the photo below stepped along a soft deposit of sand left by a recent flood. The raccoon moved from the upper right to the lower left, leaving tracks in the sequence right rear, left front, left rear, right front. The difference between the wider but tighter rear track and the narrower, more spreading front track is easily seen in the set of prints at the upper right. Raccoons habitually work the edges of streams and ponds where they find tasty shellfish, frogs, crayfish and other invertebrates. The pattern of alternating sets of hind and front tracks from opposite sides tells us the animal was moving at a pace-walk.

Mink are also in the habit of travelling along the margins of water bodies. The animal that made the tracks in the next photo was moving from right to left at a lope, and the track sequence is right front, right rear, left front, left rear. Like raccoons, mink have five toes on both front and rear feet, but it’s not uncommon for the impression of the inner toe to be missing. In fact the only print in the photo that shows a clear inner toe is the left front. This track also shows the middle pad protuberances (just behind the toes) and the heel pad (the small indentation behind the middle pad). Mink share a taste for crayfish, frogs and invertebrates with raccoons, and occasionally catch small fish. They’re adaptable predators and may also hunt for small mammals on the surrounding land.

The mink’s larger relative, the river otter, also leaves its tracks along the edges of ponds and streams, but for this creature it’s mainly a matter of convenient travel between feeding areas. I found the tracks in the photo below on the inside of a bend in a stream where an otter had taken a short cut across a large sandbar. The sequence of tracks is the same as that of the mink tracks in the previous image, and the family resemblance–both mink and otters are mustelids–can be seen in spite of the different substrates. Otters are more aquatic than mink and capture most of their food in the water.

When they’re not foraging in the water otters spend their time on conveniently accessed sites near the water. They roll on soft surfaces like grass and forest duff to clean and dry their fur, and they socialize with other members of their family group. They also leave notices in the form of scat to non-resident otters that the territory is occupied. The otter scat in the photo below contains crayfish shell fragments, but it’s also common to find scats containing fish scales and bones, or the slimy remains of frogs. Otters often use latrines where scat of various ages and contents can be found.

The beaver is another very aquatic mammal. In the photo below you see two beaver tracks, a right front (above) and a right rear (below), both facing toward the right. In the front track the four toes show clearly and the two heel pads appear as elongated grooves because the foot slipped in the mud. In the bottom part of the frame the three outer toes of the hind print show clearly but the two inner toes are obscured by the front print. As is often the case, the webbing of the hind foot doesn’t show. The size difference between the front and rear tracks is striking and helps us to understand why beavers are such strong swimmers. Beavers feed on the leaves, bark, and stems of woody plants year-round, but during the growing season the diet also includes aquatic plants, cattails, sedges, and forbs. Their tracks usually lead between the water and foraging sites on land, and signs of branches being dragged into the water are common.

Smaller–but just as well adapted to life in water–is the muskrat. Like the beaver, the muskrat has rear feet that are much larger than the front. In the photo below, the track farthest to the left is the right rear, and just to its right you see the right front. On the right side of the frame the left rear lies below the left front. Notice that the small inside toes of the front feet made impressions in both of the front prints. The muskrat’s front feet, like those of the beaver, are adapted for handling food items and building materials rather than for swimming.

If you wander along shorelines you may find muskrat latrines. These sites are usually located on logs or rocks that lie in the water but protrude above water level. In the next photo you can see a rock decorated with scat of varying ages, deposited as an announcement that the territory is occupied. Although muskrats occasionally consume animal foods they are primarily plant eaters, and their scats usually contain fibrous material.

This is just a sampling of some of the wonders to be found along the margins of lakes, streams, and marshes. There’s always something to be discovered, so next time you’re out and about, take a detour to check a stream edge or a muddy shoreline. Better yet–if you don’t mind some wading–try a stream walk. It could be just the thing on a summer day.

Cottontail Rabbits

Familiar animals can be just as interesting as less common ones, and the cottontail rabbit ranks as one of our most familiar–and interesting–creatures. In the photo below (direction of travel from right to left) we see it’s characteristic Y-shaped bounding pattern: two rear tracks even with each other and widely spaced, and two front tracks behind the rear ones, more narrowly spaced with one leading the other. The right front print (the first foot to come down) lies at the right side of the photo and the left front print (the second foot to come down) lies to its left. Farther to the left you see the rear prints which form the diverging branches of the Y. I found these tracks on a highly developed barrier island on the New Jersey coast, probably not a place you would expect to find cottontails. But these animals manage to survive and flourish not just in rural and undeveloped areas but also in city parks, suburban communities, and busy commercial zones.

Although the pattern shown above is very common, it’s not the only four-print arrangement you’ll see. Sometimes a rabbit’s front feet come down together, and when this happens the prints are even with each other and pressed tightly together. Bounding squirrels make groups similar to those of rabbits, but the spacing of the front tracks is different. Whether the front prints are even with each other (the most common arrangement) or whether one leads the other, there is almost always a gap between the two prints. In the photo below the rabbit tracks are in the lower left and the squirrel tracks are at the upper right.

The tracks in the photo below were made by a cottontail bounding in deep snow (direction of travel from bottom to top), and the toes are splayed out in both front and rear tracks. Tracks like these are sometimes mistaken for snowshoe hare tracks because of their larger size.

The feet of both cottontails and snowshoe hares can spread when increased support is needed, but there’s a drastic difference between the two animals. The maximum width of a cottontail’s hind print is about 2 1/2 inches, while a snowshoe hare’s rear track can reach a width of more than 5 inches. The photo below shows a rabbit’s rear foot (seen from the bottom) in a splayed position. Note that the rear foot has only four toes.

In the photo above you can see the thick fur which covers the bottom of the rear foot of the cottontail, and the front foot is just as furry. This is why the outlines of the toes in rabbit tracks are blurry, especially in snow. The next photo shows the right front print of a cottontail (facing to the right) in mud that had dried to a perfect consistency for recording fine details. The toes are visible but not sharply defined, and the texture of the fur can be seen in and around the toe impressions. This photo also shows all five toes clearly–yes, there are five toes on the front foot of the rabbit. But counting toes can be difficult because there are also some pads which look like toes.

To help sort this out I’ve marked the toes and two of the pads in the next photo. The innermost toe is marked Toe 1, following the convention of numbering from the inside of the foot. It’s smaller than the others and often fails to register in tracks. The other four toes are larger and tipped with substantial claws, and the toe arrangement as a whole is asymmetrical.

If you’ve ever had a run-in with a rabbit’s foot you know that, in spite of the furry covering, the sharp claws can dig in quite effectively. Sometimes the claws are the only parts of the foot that make impressions, as in this photo of the right and left rear tracks of a rabbit in a hurry (direction of travel toward the upper right).

In addition to tracks, rabbits leave many other signs of their presence. You may find stems bitten off at an angle like the multiflora rose in the photo below. These angled cuts are characteristic of rabbit browsing and they arise from the anatomy of the rabbit’s jaws.

In the next photo you see the lower jaw of a cottontail with an added line representing a stem or twig. As it takes the stem between its upper and lower incisors, the rabbit positions the stem so that one end passes through the gap between its incisors and its molars. This biting technique results in an angled cut. Deer don’t have upper incisors so instead of making a clean bite, a deer grasps the stem between its lower incisors and its horny upper palate and pulls or jerks to make a rough break.

Cottontails also feed on the bark of young trees and shrubs. Their chews have a rough appearance, with bites penetrating to varying depths, as in the staghorn sumac stem shown below. Chews made by other bark feeders (beavers, porcupines, voles, and occasionally squirrels) are much neater and more consistent in depth of penetration.

Whether it’s bark, twigs, or buds, a rabbit has to ingest a lot of fiber to get at the nutritious living cells in the cambium or in the tiny leaf initials inside buds. The animals boost the nutrition they get from their food by processing it twice. After passing through most of the digestive system, waste is diverted to the caecum where it is fermented to produce additional nutrients. This material is eliminated, usually at night, as clusters of soft globs called caecotropes. We seldom see this kind of fecal matter because the rabbit eats it immediately. After passing through the digestive system again, the waste is eliminated as pellets like the ones in the next photo.

These pellets are dry and fibrous, and are normally scattered irregularly where rabbits feed and move about. Unlike the rounded cylindrical pellets of deer, rabbit pellets are shaped like slightly flattened spheres. Cottontails are now shifting to their summer diet of grasses, forbs, and flowers, but the final result will be pellets similar to those produced from woody food.

The cottontail rabbit is a thoroughly interesting creature with some impressive tools for survival. By observing its tracks and trails as well as chews, scat, and other sign, we can appreciate a creature that is beautifully adapted to its environment.

Beavers at Work

Some animals live among us almost undetected, and others leave evidence that is obvious and long-lasting. Beavers are a good example of the latter, leaving signs of feeding on woody plants and creating dams and lodges that may last for years. The featured photo shows a lodge made of mud and sticks and surrounded by protective moat of water. What we can’t see is the underwater entrance which leads upward to a dry, multi-level living area. In the upper left quadrant of the photo there’s a dam, seen from the upstream side. The freshly peeled sticks that decorate the lodge and the dam, and the water lapping right up to the top of the dam, indicate that there were beavers in residence when the photo was taken.

Beaver dams can be impressive structures. The next photo is a view of a dam from the downstream side. The heaped up sticks conceal inner layers of mud interlaced with more sticks. Beavers react to the sound of flowing water, and any leaks are plugged with mud and more sticks. As long as the dam is maintained, the water level stays high enough to keep the lodge (not seen in this photo) secure in its watery surroundings.

A little exploration around the edges of an active beaver colony will turn up additional signs. When whole trees are removed the only thing left may be a chewed stump like the one below. Wood chips scattered nearby show where branches were removed and the trunk was sectioned and dragged off.

Some logs may remain where they fell, but they seldom go unused. One such log is shown in the next photo, and it’s a nice demonstration of the beaver’s chewing technique. The horizontal row of small cuts along the edge of the bark shows where the upper incisors were anchored. The vertical grooves in the lower peeled part of the log were made as the lower incisors were drawn up toward the anchored upper incisors. And why all this chewing? To get at the living cambium cells, located between the wood and the outer bark. Although there are exceptional cases where other food is available (such as the rhizomes of water lilies), most beavers depend on the cambium of woody plants for survival when leafy vegetation isn’t available.

Many dams and lodges are used for several years in succession, and newly added sticks and branches stand out against the mud and older sticks. The residents of a beaver colony also create deliberate messages indicating their claim to the location. In the photo below you see mud that was dredged up from the bottom and deposited on top of a grass hummock to create a scent mound. Whatever is handy at the edge of the pond, whether it be mud, muck, or rotting vegetation, can be dredged up and carried to the shore to make a pile. The final touch comes when the beaver drags its anus over the mound and deposits urine and secretions from its anal glands and castor sacs. The smell is not unpleasant, but it’s hard to describe. It reminds me of a horse barn, but it has also been compared to musk, human sweat, cheese, fruit, leather, birch beer, or some combination thereof. Scent mounds are most often created in spring, and the distinctive odor can persist for weeks.

If ponds and small streams aren’t available, beavers take up residence in creeks and rivers. But these habitats are subject to regular flooding, and the volume of flow during high water would destroy dams and lodges, so river beavers make their homes in the stream banks, digging underwater entrances and excavating living spaces above water level. If beavers are living in a river you may find peeled sticks, cut stumps, and scent mounds along the riparian margin. Another good clue is tracks in silty or muddy stream margins.

The tracks in the photo below were made by a beaver walking from lower right to upper left. The large hind prints make a wide zig-zag, starting with the left rear in the lower right corner. As in other four-footed walking trails, there are two footfalls–a front and a rear from the same side–at each zig or zag, but the front prints are mostly covered by the larger rear prints. All, that is, except for the two tracks at the upper left. The larger print above is the right rear, and next to it, just below, is the much smaller right front. Some of the rear prints may remind you of the tracks of a large bird. That’s because beavers often touch down lightly or not at all with their two inside toes, so the outer three toes make the most prominent impressions. But the heel marks behind the toes (as well as the wide palm areas at the bases of the toes) tell us it was a beaver, not a bird.

Whether they’re pond beavers or river beavers, at some point the animals will have exhausted the local food supply and will be forced to relocate to a better situation. If dams and lodges are not maintained water levels will fall. Chewed stumps and peeled sticks will weather to dull gray. Even though they aren’t fresh, these signs will persist and provide clues to the past presence of beavers.

But an abandoned beaver pond can offer its own discoveries. The amount of material amassed to form a dam or a lodge can only be appreciated after the water is drained. The exposed mud can be a great place to find the tracks of other creatures. And old beaver ponds provide great opportunities to find beaver scat. The animals normally defecate in the water, so seeing fresh scat is rare. But once the water has drained out, scats may be left on the sloping inner sides of the dam and the perimeter of the pond. Beaver scat is oblong and can be anywhere from 1/2 to 1 1/4 inches in length. The fibrous content is easy to see in the photo below–a beaver needs to chew through lots of bark and wood to get enough nutrition.

I never get tired of visiting beaver sites, because there’s always something new to discover. Whether it’s the prodigious size of a felled tree, the clever way the animals engineer channels to make transporting logs easier, or a muddy stream margin decorated with the tracks of beavers commuting to and from work, it’s always fascinating. Beavers lead complex lives and show great ingenuity in dealing with their surroundings, and the signs they leave can give us a window into their cleverness and adaptability.

Checking Out Fox Dens

It’s late summer, and fox pups that started life back in late March or early April as small, helpless balls of fluff have grown into sleek coated adolescents. At about five weeks they emerged from the den, and for the next month or so they spent their time near the burrow, developing their strength and coordination through play and mock fighting. They were guarded by parents or females from the previous year’s litter, and at the first sign of danger they dashed to safety below ground. Later they began to accompany their parents on explorations and hunting forays. But even when the kits were quite mobile, the family may have continued to rely on the den as an emergency refuge. I don’t like to disturb fox families until they no longer need that safe haven, so I always wait until late summer or fall before I investigate a fox den.

Fox dens are found in settings ranging from dense cover to open, exposed areas. The den in the photo above was near a house and driveway, but it was well concealed by shrubby cover and wasn’t obvious until I got up close. In the next photo you see the other extreme–a den dug under an old pallet that was located near a highway intersection and lacked any concealing cover. Both of these are red fox dens.

A closer shot of the den in the photo above shows several entrance holes, a throw mound (at the lower right), and tracks in the loose dirt. These photos were taken during the excavation phase, and by mid-spring the den site–and the antics of the pups–were hidden by a screen of grasses and forbs.

Cemeteries are another popular spot for red fox dens. The one in the photo below was behind a grave stone and shaded by a mulberry tree. The black specks you see scattered on the dirt and grass are mulberries.

The mulberry tree was a handy source of food, and fox scat containing mulberry seeds decorated some of the grave markers.

The next photo shows a gray fox den. There wasn’t any concealing cover around the entrance, but getting to it involved a thirty-minute hike and a climb up a steep slope. Gray fox dens are usually farther from human habitation and harder to get to than red fox dens.

There are other medium-sized mammals that use burrows, and distinguishing among them can be tricky. Tracks or direct observation of the residents are the best clues, but failing that we have to rely on other features. One helpful characteristic is the size of the entrance hole: fox dens normally have openings between 6 and 12 inches across, while coyote dens can be twice that. (And like gray fox dens, coyote burrows are not generally found in places with a lot of human activity.) Woodchuck dens have entry hole sizes on the small end of the fox range and can be difficult to separate from fox dens. The most striking difference between the two is the condition of the soil around the entrance. Woodchucks use dens throughout the growing season, and the throw mound, a spreading apron of fresh dirt like the one in the photo below, shows signs of disturbance as long as the resident is active. But unlike fox dens, woodchuck dens show little disturbance of the surrounding soil. These summer lodgings may be abandoned for burrows in more sheltered areas when it’s time for hibernation, and once that happens the soil near the summer den loses its disturbed look.

Fox dens can also have throw mounds (as in the third photo), but they usually have a more trampled look due to the comings and goings of the parents. Once the kits emerge and begin playing, their antics create an area of flattened soil and plant growth much larger than the original disturbance. Active fox dens may also have scat and the bones of small animals nearby. Occasionally, as in the photo below (also in a cemetery), other kinds of toys show what the pups have been playing with.

By the time fall comes the young foxes no longer need an underground refuge, and the area around the den becomes weathered and undisturbed. Unused dens are difficult to assign to an owner and may, in fact, have different residents in the next birthing season. During the winter foxes begin to investigate den sites, and an alert tracker may notice the telltale tracks of a pair of foxes making frequent trips to certain locations. Once the snow is gone favored sites show signs of activity like those shown in the second and third photos.