Furry Feet

It’s been awhile since I’ve put out a new post, due to some medical issues I’ve been having. But things have improved, so I’m happy to be back in action.

Going barefoot, or more accurately, bare-toed, is quite popular among wild mammals. The tops and sides of the feet are typically furred, but most mammals in our region touch the ground with bare skin. This applies to both toe pads and middle pads, and sometimes the heel area as well. In the photo of red squirrel bounding tracks below (direction of travel toward the top) the hind tracks are set more widely above the front tracks. The clean borders and smoothly compacted interiors of the toe and middle pads–and the heel pads in the front prints–are unmistakable evidence of bare skin.

But there are a few inhabitants of our region that buck the trend. The feet of rabbits and hares, for example, are thickly furred, and this fur has a blurring effect on the impressions of the toes and pads. In the photo below you see the tracks of a cottontail rabbit bounding toward the left. The toes are visible, but they lack the clear outlines and smoothly compacted interiors seen in the squirrel tracks shown above.

The fisher is another animal with thickly furred feet. The tracks in the photo below were made by a fisher loping from lower left to upper right, and the toes and middle pads are almost completely muffled by the fur.

In the preceding photo the snow was cold and dry, but if the snow is wetter the toes may be more recognizable. The next photo shows tracks made by a loping fisher traveling from lower right to upper left. In the warmer and more easily compacted snow the toes made visible impressions, but the blurring effect of the fur can still be seen in the middle pad and heel areas.

River otters share many features with fishers but differ in having unfurred toes, middle pads, and heel areas. In the next photo there’s a collection of otter tracks (as well as a few dog tracks). The otter loping pattern in the middle and lower part of the photo (direction of travel from lower right to upper left) is the important part, but it’s a little confusing so bear with me and I’ll try to sort things out. Starting at the lower right the sequence of the otter loping tracks is right front, left front, right rear, left rear. There are a few more otter tracks in the upper area of the photo, and in between the two otter front prints there are two dog tracks, a front with a rear just to its left, oriented to the right. The otter prints in the loping array, especially the left hind print at the far left, show the clearly outlined toe and middle pads and the smooth heel areas characteristic of bare skin. And incidentally, the rear dog print also has cleanly outlined toe impressions made by toes of bare skin.

We have two species of foxes, one with furred feet and the other without. The unfurred feet belong to the gray fox. In the cluster of gray fox prints in the next photo the rightmost and leftmost tracks were made by front feet, and the clearer of the two tracks in the middle was made by a rear foot. The distinct margins bordering the toes of both front and hind prints, and the middle pad of the front track on the right, are evidence of skin covered pads.

Red foxes have thick fur covering almost all of the undersides of the feet, and the blurring effect can be seen in the next photo (rear print on the left, front print on the right, and direction of travel toward the left). But here things get a little complicated. I said the feet were “almost” completely covered because there are two places that don’t grow fur. In the middle pad of the red fox’s front foot there’s a curved ridge, sometimes called the bar, which is unfurred–it can be seen in the front track at the lower right in the photo. There is also a small, hairless oval near the tip of each toe on both the front and the hind feet. In the photo these bare regions show as dark spots in the tips of the toe impressions.

Although tracks in snow usually reveal the presence of fur, mud is better for rendering the fine details. In the next photo you see the front print of a red fox oriented to the right. The fur pressed into the mud shows as thin striations in the toe and middle pads. The unfurred parts of the foot, the bar in the middle pad and the small hairless ovals in the toes, interrupt the striated texture left by the hair.

Felines are characterized by bare toes and middle pads, so their tracks have the cleanly outlined pads of other animals with unfurred feet. But bobcats have separate, interesting feature connected with fur. The fur covering the tops and sides of the feet is very thick, especially in winter. When snow conditions are right this fur pushes into the snow around the outside of the track, creating what’s called a hair halo. In the photo below of a bobcat’s right front print (oriented toward the right) the hair halo shows as a sloping bevel outside the toes.

Hair halos aren’t generally seen in house cats–or canines for that matter. In the next photo you see a coyote front print bordered by abrupt walls rather than a sloping bevel. The inclines at the front and back of the track are entry and exit disturbances, and the grains of snow scattered over the track are due to a light snowfall which must have ended just after the track was made. And as a side note, only bare toes could have made the very distinct and cleanly outlined toe impressions you see in the photo.

Do furred feet help in survival? Probably, since thicker fur over the whole body develops as winter approaches. Are furred feet essential to survival? Probably not, since only some mammals have this trait. For the tracker, it’s enough to recognize furred feet as an interesting characteristic and to appreciate the effect fur has on tracks of some animals.

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.

Dog Tracks: Common But Not Always Commonplace

Our familiar companion animals leave their tracks everywhere, and we see them so often we may find ourselves giving them only superficial attention. But if we look more carefully we may be surprised at how much they have to teach us. Dog tracks can show us the traits that are shared among dogs and their wild relatives, and they can also reveal the remarkable range of variation that could only belong to a long-domesticated animal.

The photo below (direction of travel toward the top) illustrates a number of core features that are shared by dogs, foxes, coyotes and wolves. The front track (lower left) is larger than the rear track (upper right). Both front and rear tracks have four toe impressions, and a single smooth middle pad. The tracks are symmetrical, meaning that if an individual track were divided down the middle and the right half were folded over onto the left half the two halves would superimpose almost perfectly.

There are a few additional traits in these prints that are shared among some–but not all–dogs and their wild relatives: The overall outlines of both front and rear prints are oval, the toes are held tightly together, and the claw marks point straight ahead. The larger ridges between the toes and the middle pad form the celebrated canine X, and the center of each track is occupied by a rounded dome or plateau. But although the tracks are roughly coyote sized, we see several additional traits that indicate dog rather than coyote. The claw marks are wide and deep, the middle pad of the front print is large compared to overall size of the track, and its trailing edge is rounded.

A comparison between the dog tracks above and the coyote tracks in the next photo will highlight the differences. (The coyote front print is below and a little ahead of the rear, and the direction of travel is to the right.) Like the dog tracks, the coyote prints are compact, with oval or egg-shaped outlines and tightly held toes. But some of the claw marks are missing, and those that are visible are delicate rather than robust. The middle pad of the front track is relatively small, and its trailing edge is concave.

The dog front track shown in the next photo is even more of a departure from the two preceding illustrations. Instead of being oval the print is round in outline, and the toes, especially the inner and outer ones, are widely spread and angled to the sides. The upper arms of the canine X seem to have been pressed apart, and there’s a curved ridge instead of a dome at the center of the track. Like the previous dog tracks, the print in the photo below has wide claw marks and a large middle pad. A track like this is easily recognizable as dog.

The photo below shows another dog track variant. The overall outline is rounded–in fact the print is a little wider than it is long. In contrast to the triangular middle pads of the previous dog tracks, this print has a trapezoidal middle pad with a broad leading edge. Instead of an X the internal ridges form a distorted H shape, and there’s a horizontal ridge rather than a dome in the center of the track. The absence of claw marks is unusual for a dog print. This is the kind of track that might be mistaken for a feline–a house cat, bobcat, or even a cougar depending on the size of the print. But there are clues that indicate dog, and they become obvious if we look at a true feline print.

The photo below shows the right front track of a bobcat. If we test for symmetry using the folding test described in the second paragraph, we can see that the dog track above is symmetrical and the bobcat track below is asymmetrical. There’s a leading toe (second from left) and a trailing toe (rightmost) in the bobcat print, and the middle pad is canted to the right. Instead of the canine X we see an angular C-shaped ridge that is also canted to the right. Another important feline trait is the contrast in size between the large middle pad and the small toes.

Wild canines have a way of placing their feet quietly, without tension or unnecessary movement. Dogs, on the other had, often express slight jiggles or shifts in the movements of their legs, and the difference can sometimes be seen in their tracks. In the photo below you see two dog tracks (in the upper left, the front a little below and behind the rear) and two coyote tracks (the front in the upper right corner and the rear just below the two dog tracks). Displaced bits of snow lie inside and around the edges of the dog tracks, while the coyote tracks have mostly smooth floors and margins. The best way to see the action that produces this kind of difference is to watch a dog walking or trotting directly away from you. You may see subtle shifts in the body or slight wobbles as the legs contact the ground.

Dogs lead easy lives compared to wild animals, and this often shows in the trails they make. The tracks of three creatures can be seen in the photo below. A dog meandered from top left to bottom right, leaving two partly superimposed tracks in the upper left corner, two tracks close together midway down the left side, and two more widely separated tracks at the lower right. A coyote trotted from top to bottom leaving smaller direct register prints, one between the dog tracks in the upper left and another at the lower left. And my boot tracks can be seen to the right of the dog tracks.

Wild creatures cannot afford to waste energy. There are exceptions: young animals play, and during mating season mature animals can make some wacky moves. But the business of survival demands efficiency of movement. Dogs, on the other hand, can expend energy without the pressure of finding their next meal. The coyote in the photo above had a destination in mind and moved with purpose. The dog was well fed and carefree, and counted on finding food and shelter when it got home. And I was focused on the story those tracks told about the lives of dogs and and their wild relatives.

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.