Zig-Zags

In past posts I’ve used the term zig-zag to describe certain track patterns. In this article I’d like to delve more deeply into how zig-zags arise and what they can tell us about the animals that make them. When we humans walk in a relaxed, natural manner we place our feet in a zig-zag pattern because each foot falls to its own side of the line made by our moving center of gravity, the center line of the trail. It’s easy to verify this: Just walk naturally in snow or mud or on a dry surface with wet feet and then look at your tracks. The same logic applies to birds, so we often see patterns like the one in the next photo, made by a turkey walking from left to right. Each print angles inward, which helps to distinguish right from left. The sequence, starting at the left, is right, left, right, left, right.

Two legged zig-zags are pretty straightforward, but four-footed animals also create zig-zags, and it’s not as easy to understand how a four-footed animal can do that. Watching animals helps, but it’s hard to follow foot placement when animals are moving in real time. Fortunately for us twenty-first century trackers, there’s a tool that can bridge the gap–the internet. So let’s take a look at a video of a horse. If you click on this link: Bing Videos, then click on horses walking youtube and start the video, you’ll see a horse walking in slow motion. Notice that as each front foot leaves the ground the rear foot on the same side comes down in the spot just vacated by the front foot. The video doesn’t show the pattern on the ground, but it’s easy to see how the horse leaves a series of double impressions, each one a front track overlaid by a rear track. And since the feet on each side fall to their own side of the center line, the overall pattern is a zig-zag. The trail in the next photo, made by a deer walking from bottom to top, is a good example of a zig-zag made by a four-footed animal.

But all zig-zags aren’t the same. The physical characteristics of animals vary, and this affects the kinds of patterns they leave when they walk. There are also different types of walks, with differing relative placement of the front and rear tracks. In the photo above the walk is an almost perfect direct register gait, meaning that the rear feet fell almost exactly on top of the corresponding front tracks. The next photo shows tracks made by a woodchuck walking from lower left to upper right (and just below the second impression, tracks of a squirrel bounding toward the bottom). The trail is more variable but the tracks are mostly in indirect register, meaning that the rear tracks fell partly but not completely on top of the corresponding front tracks. Starting at the lower left the track sequence for the woodchuck is: right rear on right front, left rear on left front, right rear, right front, left rear on left front. Even in this more irregular trail the zig-zag is apparent.

The width of the zig-zag, known among trackers as trail width, varies from one species of animal to another. To measure trail width, find a relatively straight part of the trail and imagine or draw out two parallel straight lines that just touch the outsides of the alternate sets of tracks. Then measure the perpendicular distance between the lines. This is diagrammed in the next photo of the indirect register track pattern made by a walking opossum heading toward the upper right.

In the next photo you see a trail made by a gray fox walking from right to left. The trail has a different look from the opossum and woodchuck trails, both because of its narrower width and also because the fox’s step lengths are longer. But the zig-zag is still apparent. Trail widths, combined with step length, can be helpful in identification, since chunky animals like woodchucks and possums make wider trails and take shorter steps than slimmer, longer-legged animals do. And trail widths are especially important when you’re considering animals with similar step lengths. For example, trail widths for a walking coyote are generally between 4 and 5 inches while trail widths for deer moving at a walk range from 5 to 10 inches. Even when the tracks are degraded or obscured by collapsing snow it’s usually possible to differentiate between a coyote trail and a deer trail.

Animals find it harder to move in deep snow, but when they’re walking their trails still show the zig-zag pattern. In the photo below a red fox walked from bottom to top leaving a zig-zag arrangement of deep holes in the snow.

All of the gaits discussed above (and the one the horse was doing in the video) fit into what I call the regular walk–also called the diagonal walk in the tracking literature. But that’s not the only kind of walk animals can do. A common variant is the overstep walk. To see a dog doing the overstep walk click on this link: Bing Videos and then click on dog gaits youtube and start the video. The recording shows a dog walking at actual speed followed by the same sequence in slow motion. If you keep your eye on the spot just vacated by a front foot you’ll see the corresponding rear foot come down a little past it. (This video also does a nice job with the amble, equivalent to the pace-walk of the raccoon, and the trot.)

The interesting thing about the overstep walk is that the pattern of tracks on the ground also makes a zig-zag, but the points of the zig-zag consist of sets of two prints, front and rear from the same side, rather than the impressions of two superimposed tracks. In the next photo you see an overstep pattern made by a house cat moving from lower right to upper left. Because a cat’s front tracks are wider and shorter than the rear ones we can see that in each set the front track is behind the rear. The sequence, starting at the lower right, is: right front, right rear, left front, left rear, right front, right rear. Among animals that are habitual walkers, overstep walks are common.

Another variation you’ll come across is the understep walk. The next photo shows the trail of an opossum doing an understep walk, heading from the lower left to the upper right. Again, the prints are arranged in sets of two, each set the front and rear from the same side. In each pair the hand-like hind track, with its thumb pointing inward, lies behind the front track with its more evenly spread toes.

We sometimes find zig-zag walking patterns in the trails of animals that aren’t habitual walkers. Fishers move mostly in bounds or lopes, but they walk when extra caution is needed or when the footing isn’t secure. The trail in the photo below was made by a fisher walking, mostly in direct register, from lower right to upper left.

Walking trails are less common for minks than for fishers, and for minks it seems to be mostly about the animal’s dislike of unstable surfaces. In the next photo a mink walked from right to left through mud (looking pretty dry in the photo but probably much wetter and slipperier when the tracks were made), leaving sets of paired tracks. But which walk is this, overstep or understep? We can tell because the middle toe in the mink’s hind print usually angles a little to the outside. So the sequence, starting at the right, is: left rear, left front, right rear, right front, left rear, left front, right rear, right front, and this is an understep walk.

White-footed mice are even less likely to walk than minks, but the next image attests to the fact that they do it on rare occasions. A white-footed mouse walked from bottom to top, leaving sets of paired tracks. The four-toed front prints lie behind the five-toed rear prints in each set, so the mouse was doing an overstep walk. The trail both before and after the walking part was on drier footing with normal mouse bounding patterns, so it was the wet mud that made the mouse shift to a walk.

Many animals get around mostly at a walk, and zig-zags abound in the tracking world. The details of the patterns can tell us a lot about the nature of the track maker. But the sight of a zig-zag for an animal whose default gait is not the walk is an even more compelling call to investigate. In addition to their help in species identification, zig-zags can tell us how animals interact with each other and with their surroundings. In this post we’ve only made a start. There are other kinds of zig-zags, and even patterns that look like zig-zags but aren’t. I’ll keep these topics for a future article. In the meantime, follow the zig-zags wherever they lead.

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.

Gray Fox Affairs

It’s been a strange winter. In my neck of the woods we had some significant snow early in the season, but no big storms since then. Temperatures have been up and down (more up than down), and with all the melting, the snow we do have has consolidated into a dense, icy layer. Much of the time the conditions have been terrible for tracking, but every once in a while something wonderful has happened: warmth and liquid precipitation have been followed by dropping temperatures and a change from rain to snow. When this happens, snow that falls while the air is still relatively warm becomes bonded to the crust. As the temperature drops and additional snow falls, it forms a soft layer on top. The result is a non-slip and easily navigable surface that is a perfect medium for recording tracks.

A few days ago I encountered just such conditions: an icy base covered by a thin layer of soft snow. I was in an extensive natural area, and both the forest road I was following and the surrounding landscape offered beautiful tracking conditions. Animals of all sorts had been moving easily over the snow, and there were tracks everywhere. I found myself following the trail of a gray fox. The animal went for quite a distance at an easy trot, but then it did something that was quite puzzling.

The photo below shows the fox trail as it goes from upper left to lower right. (You can also see a coyote trail to the right of the fox trail, and a mountain bike trail to the right of that.) As it entered the frame the fox was walking (the first three tracks at the upper left). In the next section (between the last walking step and the edge of the tree shadow) the pattern was very different, and following that the trail looks unlike either of the previous sections. I wanted to know what was going on and why the middle section looked so different.

In a situation like this the first thing to do is identify each track. The zig-zag of the walking section helps us to tell right and left, and the fact that the front foot is larger than the hind foot distinguishes front from rear. The next photo shows a gray fox front print on the left and a gray fox rear print on the right. You can see the difference in overall size and also the difference in the sizes of the middle pads.

The photo below shows just the puzzling middle section, and if you compare photos you’ll see that the front and rear in the photo above are actually the first two tracks in the middle section. It’s pretty clear that the first four prints in the photo below are left front, left rear, right front, right rear. After that it gets harder. The track just above the right rear is smaller than the one to its right, so those two prints must be left hind, left front. Three tracks from the right side come next, and it looks to me like the sequence is right rear, right front, right rear. The final two before the tree shadow are the left front and the left rear, and at the edge of the tree shadow there’s a right front with a right rear partially superimposed on it.

In the next photo I’ve added labels showing my take on right/left and front/rear. If we start at the beginning of the whole sequence, the animal was trotting (those tracks aren’t seen in the photos) and then slowed down to a walk (the first three tracks in the distance shot). The next section shows that the fox slowed even more to an overstep walk (the first four prints in the photo below), then slowed even more to an understep walk. There’s an extra right hind that’s puzzling, but I’m guessing the fox just repositioned its right hind foot. Then the overstep walk reappears after which the fox picked up the trot (the two impressions at the lower right in the first photo). Notice that the step lengths in the overstep part are shorter than the regular walk steps that preceeded them, and the step lengths in the understep part are shorter yet.

That analysis was rather involved, but it leads to a picture of what the fox actually did. As it trotted along something it detected made it slow down, first to a walk and then almost but not quite to a standstill. It was probably sniffing and listening intently as it moved very slowly. Once the animal concluded that it was okay to move on, it resumed its journey at a trot. It’s impossible, without more evidence, to know what caused the fox to react the way it did. It may have been a threat, but it could also have been something that interested it for a different reason. It is, after all, mating season for wild canines.

And the fox I was following was definitely tuned in to mating season. Farther on I found a spot (shown in the next photo) where the animal had detoured to urinate on a small spruce branch. If you look in the center of the frame you’ll see a squiggle of urine that runs horizontally from the upper edge of the spruce branch. Because the urine wasn’t squirted out the side of the tracks we know this was a female. She would have lifter one hind leg forward and supported herself on the other hind leg (plus two front legs) as she urinated. The relative depths of the tracks tell us that the supporting rear foot was the left. Its track is in the prominent double impression above and to the left of the urine.

I’ll never know what made the gray fox slow down and leave the pattern discussed in the beginning of this article. It could have been a threat–there was certainly a coyote in the neighborhood, or it could have been the mountain biker. A fisher (whose tracks I also found on that day) would have made the gray fox nervous. And there were red fox trails as well. But the trail shown in the photos above doesn’t suggest alarm so much as cautious interest. The fox didn’t change direction but just continued on. Was it another gray fox, one she was familiar with, or one she had mated with in a previous year? We have a small part of the whole story, and we can only speculate about the rest, but it’s fascinating just as it is.

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.

Mud, Glorious Mud!

Unlike many people I know, I’m always sad to see the snow disappear for good. But as soon as I think of what comes next–mud season–I get excited all over again. The transition between the seasons is highlighted in the photo below. A gray fox had stepped in some mud and then left it’s muddy footprints on the snow as it walked from left to right. Each deposit of mud is made by first the front and then the rear feet from the same side, and the zig-zag pattern of the walk shows nicely.

Once the snow is gone, ordinary mud takes its place as a medium for recording tracks. In the next photo an opossum stepped in some mud at the edge of a puddle, leaving a collection of both complete and partially obscured prints oriented toward the left. At the upper left there’s a right front track with a right rear track just behind it. Farther to the right you can see part of another right front track. In the lower right corner there’s a nearly complete left front print and just the suggestion of a left rear behind it. The front prints show the five widely spread toes that are characteristic of the opossum. In the front print at the upper left the segments of the semicircular middle pad are especially clear. The rear track in the upper center shows the strikingly different form of the opossum’s rear foot: a thumb-like inner toe pointing inward and four additional toes close together and pointing outward.

One of the nice things about mud is that it can record the presence of animals that we don’t encounter during the cold season. A spotted salamander (or maybe two of them) walked through the mud in the next photo. These amphibians hibernate in winter and come out in early spring, so mud season is a good time to look for their tacks. There are two trips: one going from the left side toward the upper right and the other proceeding from left to right along the lower part of the frame. Each trail consists of a central drag mark made by the tail and a sequence of front and rear prints on each side. The patterns of the prints tell us that the animal(s) were moving at an understep walk, with each rear foot touching down just behind the front foot from the same side. In the lower trail you can see the difference between the smaller, four-toed front prints and the larger, five-toed hind ones.

The tracks in the preceding photos are pretty obvious, but it’s not always easy to spot tracks in mud. In the photo below there’s a patch of shiny mud in the center of the shot, and on the right side of that patch there are some tire tread marks. If you look on the left side of the same shiny mud toward the top you’ll see a red fox track. The animal was moving from top to bottom, and because there weren’t many muddy spots the print in the photo was the only one I could find.

The close-up below shows the same track, but in this view it’s oriented in the opposite direction, toward the top. The central mound typical of canine tracks can be seen, and the marks made by the hair on the underside of the foot show clearly. There’s even a partial impression of the bar in the middle pad.

Woodchucks, like salamanders, spend the winter below ground and often emerge just as mud season is beginning. The next photo shows the left rear track of a woodchuck at the upper left and a left front track at the lower right. The five clawed toes of the rear print show clearly–the middle three toes set close together and the inner and outer toes angled toward the sides. Behind the toes you can see the four segments that make up the middle pad. In the front track the four toes with their substantial claws can be seen. The subdivided middle pad of the front foot lies behind the toes, and the heel pads show as two depressions behind the middle pad. The front print has a curvature toward the inside, a trait typical of the woodchuck.

It takes a medium with a fine texture to show details of the tracks of very small animals, and what better medium than mud? In the photo below you see the tracks of a white-footed mouse bounding from lower left to upper right. The tracks are arranged in the typical rodent bounding pattern–two rear prints (in the upper right quadrant) that are widely set and almost even with each other. Behind the rear tracks, the front prints are set more narrowly and, in this case, slightly staggered rather than even with each other. Track details show beautifully, especially in the left rear (the topmost track) and the left front (farthest to the left). If we compare these tracks with the woodchuck tracks above we see the rodent family resemblance, especially in the rear prints. The symmetrical mouse front tracks are more typical of other small rodents than the curved front prints of the woodchuck.

You can’t ask for a better rendition of detail than the porcupine tracks in the next photo. Porcupines have unique foot anatomy: their tough, undivided soles have a pebble-like texture that gives the animals good grip when climbing. The photo shows a left front print and, just behind it and overlapping slightly, a left rear print. The tracks are heading toward the left, and the texture of the soles shows beautifully. Because the leading edge of the rear foot touches the trailing edge of the front track, the two tracks seem to be joined together. You may be able to pick out the claw marks of the hind print along the leading edge of the sole of the front print. The four claws of the front foot made indentations at the very left, and if you look closely there are marks made by the three outer phalanges of the front foot behind the claw marks.

Once the snow melts and the weather warms, mud may not last long. Puddles may dry up and wet areas may fill in with plant growth. But mud can also appear in new places, and abundant rainfall can bring on new mud seasons long after the early one is over. As a matter of fact, I found the porcupine tracks in the photo above in the month of July. So keep an eye on the conditions of the puddles in your neighborhood, and don’t be surprised if you come across some beautiful mud when you least expect it.

Canine Romance

It may seem like the wild creatures are all hunkered down, just doing their best to stay alive and wait out the cold season. But for wild canines there’s more going on than simple survival. This is mating season for foxes and coyotes, and they’re engaging in behaviors that will eventually lead to reproduction. For human trackers the first clue to their new fixation is the increasing frequency of scent marking. In the photo below you see a splash of urine on the snow to the right of the corn stalk, left as a message to others of its kind by a gray fox.

Urine contains complicated combinations of chemicals that, to a discerning canid, reveal the identity and health status of the animal that produced it. As mating season progresses, changes in the chemical signature also indicate the animal’s readiness for mating and reproduction. Most of these messages are too subtle to be detected by people, but fox urine is an exception. The urine of both red and gray foxes takes on an increasingly musky, skunk-like odor as hormonal changes progress, and in late winter this odor is strong enough for a person to detect it a good distance away from the deposit.

The fox whose scent mark is seen in the image above was walking from the upper left to the lower right. We know that the it was a male because of the placement of the urine off to the side of the animal’s trail. The direction of travel is revealed by the appearance of the snow around the edges of the tracks. When a foot goes down into snow it pushes any movable material down into the hole. When the foot comes up and out again it often brings a little snow up with it, and this snow is dropped around the leading edge of the track as the foot moves forward above the surface. This means that the snow around the entry end of a track is relatively undisturbed while the surface at the exit end is decorated with sprays or scatterings of snow. In the photo of the gray fox trail there are four deep tracks, each one the landing place of a front and a rear from the same side, plus a light track near the third deep track. Starting at the upper left (and considering just the deep tracks), the sequence is left front + rear, right front + rear, left front + rear, right front + rear. The animal was standing on its right hind foot when it lifted its left hind leg to squirt urine at the corn stalk. The shallow print was probably made as the fox placed its right front foot lightly on the snow for balance while it was urinating. The corn stalk was a convenient object, but rocks, clumps of weeds, branches, or anything else that protruded above the snow would have done as well.

Among coyotes and foxes, early winter is the time for the establishment or reestablishment of pair bonds. Those that spent time apart after they raised a litter in the previous season usually rejoin, and unattached animals roam widely in search of potential mates. Bonded pairs establish their territory by scent marking around the edges, especially along borders shared with others of their species. Unless we are familiar with the area and the animals involved it’s hard to know whether scent marks are advertisements of availability or warnings that the territory is occupied.

During the lead-up to mating, pairs often travel together. The trails in the photo below were made by two red foxes travelling along a forest road. For at least half a mile the two trails wove back and forth, occasionally changing speed and sometimes diverging, but always coming back together. The male, with slightly larger tracks, enters the frame at a gallop, moving from the lower left to the upper right. The female is doing a side trot and her trail comes in at the lower right and leaves at the top of the frame just to the left of the male’s trail.

All of the behaviors I’ve described above help to strengthen the pair bond and propel the hormonal changes that lead up to mating. When the female’s endometrial lining begins to develop, her urine contains blood, and she leaves scent marks like the one in the photo below. She’ll soon go into heat and only then will she be receptive to the male’s advances.

The pair are both involved in the process of den selection and preparation, and pups are born about 50 (foxes) or 60 days (coyotes) after successful coupling. By initiating the early stages in the dead of winter nature insures that the pups are born in the spring when food will become increasingly abundant. Note: It’s important for trackers to avoid disturbing animals during this vulnerable time. We should not approach too closely or otherwise disturb a den site starting with the period of den preparation and continuing until the pups are no longer dependent on the den for safety.

This is, by the way, why coyote-dog hybrids aren’t as numerous as some people believe. Over thousands of years of domestication, dogs have lost the finely tuned sequence of reproductive behaviors that occurs in wild canines. Mating in dogs is no longer synchronized with the seasons, and males don’t assist in the raising of pups. Since these behaviors are genetically controlled, the offspring of matings between dogs and coyotes have disrupted patterns of behavior. The precise timing of reproduction is lost, as well as the strong pair-bonding and the dual effort from both parents (and sometimes female offspring from the previous year). For this reason the offspring of matings involving dog-coyote hybrids are unlikely to survive.

This is a great time of year to let tracking open a window into some of the underlying processes of the natural world. The mating rituals of wild canines have been shaped for success by natural selection, and this is beautifully illustrated in the behaviors we see in the tracks and trails of foxes and coyote.

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.

Separating Felines and Canines

Cats and dogs–they live among us as companion animals, and their wild relatives are a familiar presence in our landscapes. The two groups differ from most other mammals in having four toes on both front and back feet, and in both groups there’s a smooth pad (the middle pad) behind the toes. So how to tell them apart? One thing we often hear is that canine tracks show claws and feline tracks don’t, but the presence or absence of claws can’t always be relied on. Cats can extend their claws if they need more grip, and the claws of wild canines may not touch the ground because they are naturally trimmed by constant movement. Fortunately there are other features that can help us to distinguish between canines and felines. It’s a simple matter of paying attention to details, and there are plenty of them. So let’s get started.

In the photo below you see the front track of a coyote, direction of travel from bottom to top. The overall shape is elongated, the toes are relatively large compared to the middle pad, and the outer toes are tucked tightly behind the leading toes. The middle pad has a pointed forward edge. Roughly in the center, between the toes and the middle pad, there’s a dome-like area, and an X is formed by the ridges between the middle pad, the outer toes, and the two leading toes taken together. The dome and the X are the negative spaces, areas that are not pressed down by toes or pads, and their arrangement is an unmistakable indicator for members of the dog family. (By the way, note that there are no claw marks.) And another important detail: the track is symmetrical. By symmetrical I mean that the right half is almost identical to the left half.

Here’s a way to grasp the concept of symmetry. In the next illustration I’ve added a vertical line which bisects the track. If you imagine folding the right half over along the line so it lies on top of the left half, the two halves will match almost exactly. The two leading toes are almost even with each other, the inner and outer toes are of similar size and spacing, and the middle pad seems to point straight ahead.

Now let’s compare a feline print. The photo below shows the front track of a bobcat. The overall shape is more rounded, the toes are relatively small compared to the middle pad, and they’re arranged in a wide arc in front of the middle pad. The left-most toe is smaller and farther back than the others. There’s no central dome, and the ridge between the middle pad and the toes is broadly curved or C shaped. Rather than pointing straight ahead, the middle pad is canted to the left. The arrangement of toes and middle pad makes the track very asymmetrical.

To get at the asymmetry here’s the same photo with a vertical line bisecting the track. If you do the mental experiment of folding one side over onto the other as we did with the coyote print, you’ll see that the two halves don’t match. One toe leads all the others, and the smallest one lags behind the other three. The whole print seems to be canted to one side. Because it’s asymmetrical we can tell which front print this is–it’s the left. If we were looking at an isolated coyote track it wouldn’t be possible to know if it was a left or a right.

Here’s the front print of another canine, a red fox. It’s not as elongated as the coyote print, but the overall shape is still more oval than round, and the outer toes are tucked in behind the leading toes. The central dome and canine X are unmistakable, and the track is beautifully symmetrical. In this case claw imprints show as delicate punctures directly ahead of the toes. The mud was just moist enough to show the hair on the underside of the foot and the chevron in the middle pad perfectly.

Perhaps you’re thinking that I oriented the bobcat track incorrectly, and that if it were rotated a little it could be made to look more symmetrical. But the asymmetry is inescapable, no matter how you look at it. In fact, when an animal is moving at a steady gait the correct orientation of each print is dictated by the direction of travel, and the imaginary bisecting line should be parallel to the direction of travel. The next photo shows part of a sequence of tracks made by a house cat moving at an overstep walk, a gait in which the rear print lies ahead of the front print from the same side. The direction of travel is from lower right to upper left.

In the next illustration I’ve added a line which defines the direction of travel to the same photo. (Sorry about the slight wobble, just imagine it’s really straight.) With the line as a reference, you can see that each track is canted to the outside. Also notable in this photo are the shapes of the middle pads: wide and blunt on the forward edges and scalloped on the trailing edges. The C-shaped ridges between the middle pads and the toes are especially striking.

The topic of gaits brings up another criterion sometimes said to distinguish canines from felines–direct register versus indirect register. Walking felines are said to place their feet in direct register, so that the back foot comes down exactly on top of the print of the front foot from the same side. Canines are supposed to prefer the indirect register walk, placing the back foot partly, but not exactly, on top of the front from the same side. But this isn’t a make-or-break test. The next three photos illustrate this point: first you see a bobcat direct register walk, next a bobcat indirect register walk, and finally a red fox direct register walk. The fact is that both felines and canines use both direct and indirect register walks. It’s fair to say that domestic dogs and cats are more likely to walk in indirect register than wild canines, but even among wild species there’s variation, and the difference isn’t very useful for identification purposes. And of course it wouldn’t apply in other kinds of walks, such as the overstep walk in the previous illustration.

Bobcat direct register walk
Bobcat indirect register walk
Red fox direct register walk

Our other fox, the gray fox, makes tracks that are a little more catlike. In the photo below you see a rear print, below, and a front print, above. The overall shapes are round enough to be confusing, the central dome and canine X are a little different from the ones shown for the fox and the coyote, and in the hind track there appears to be a leading toe. Gray foxes have semi-retractable claws which don’t usually show in tracks, so the presence of claw marks suggests that the animal needed more stability in the wet mud.

In spite of the variations the tracks are clearly canine. Compare them to the bobcat tracks in the next photo, this time a left rear below and a right front above.

These two photos bring out some potential pitfalls in the task of differentiating canines from felines. Animal feet aren’t rigid, and toes may spread or tighten depending on the animal’s movement and the nature of the substrate. In the gray fox hind print the two leading toes gripped the soil differently, causing one to appear farther ahead than the other. The normal symmetry of canine tracks can be altered by head turns, changes of direction, or by sloping terrain. Another potential source of uncertainty is the fact the hind feet of felines are often more elongated than the front feet. In the bobcat photo above the rear track has a slightly canine appearance due to the tighter arrangement of toes and the more oblong shape.

And lastly, never underestimate the power of domestic dog tracks to create confusion. The track in the next photo was about the size and overall shape of a bobcat print, lacked claw marks, and mimicked the blunt middle pad and C-shaped ridge of a feline. Red flags were raised by the symmetrical structure and the large toes, but it was really the presence of more tracks which clinched the identity as dog. Domestic canines are incredibly variable–their tracks can be quite round and the tendency of the toes and middle pads to spread can give the negative spaces a feline appearance.

So the task of separating canines and felines is not always easy. Isolated prints are harder than more complete sequences, and weathering and distortions can make things difficult. But the more you study them, the better you’ll be at picking out the crucial features. And if one of these creatures is rare in your area, it’s a joy to find it’s tracks and know you’ve made a solid identification.