A Family Resemblance

Rodents are the most common mammals on earth, in both number of individuals and number of species. They are also the most diverse, with lifestyles that range from semiaquatic through fossorial (adapted for digging and living mostly underground), terrestrial, arboreal, and even semi-aerial (gliding flight). But don’t let that mind-boggling profusion intimidate you. In our region many of the most common rodents are members of the squirrel family, a group that is remarkably uniform in physical features. Fortunately for the tracker this uniformity extends to track details and track patterns, and familiarity with the key features will aid in the recognition of any member of the group.

In the photo below you see tracks made by a gray squirrel bounding toward the top of the photo. The five-toed rear tracks lie in the upper part of the image, and the four-toed front tracks can be seen in the lower part. Claw marks show as tiny pricks ahead of the toes of both front and rear tracks. Notice that the toe pads of the three middle toes of each hind print are lined up close together, while the inner and outer toes lie farther back and angle to the sides. Behind the toes you can see a C-shaped grouping of middle pads. The front tracks have only four toes, but again the central two point more forward while the outer and inner ones point to the sides. C-shaped arrays of middle pads sit behind the toes of the front prints, and heel pads (there are two on each foot, but it’s hard to tell in this image) are situated behind the middle pads.

Bounding is the most common gait for most members of the squirrel family, and the resulting pattern is another recognizable trait of the group. In the photo above the two rear prints are almost even with each other and are set wider and well ahead of the front ones, which are also nearly even with each other. This positioning may seem odd, but there’s a logical explanation. At each bound the animal lands on its front feet and draws its rear feet forward so they pass outside of its front legs. As the front feet lift off the rear feet touch down–ahead of the spots the front feet just left–and propel the next leap.

The next photo shows a bounding pattern made by a red squirrel, again travelling from bottom to top. There’s a striking similarity to the first image of the gray squirrel tracks, in both overall arrangement and track details. Because the substrate was softer the rear feet of the red squirrel (in the upper part of the frame) sank in deeper–notice that the whole length of each of the three middle toes registered as a narrow groove. Nevertheless the three toes are closer together and oriented more forward than the outer toes, just as they were in the gray squirrel tracks. In the front tracks of the red squirrel (in the lower part of the photo below) the claws show as grooves rather than pricks, but the overall structure is similar to the front tracks in the preceding shot. If you look at the red squirrel’s right front print (at the lower right in the photo below) you can see clear impressions of the two heel pads.

The chipmunk tracks in the next photo (again bounding toward the top) are consistent with the features we saw in the red and gray squirrel prints. In the right rear print (in the upper right quadrant) you can see that the middle toes are closely grouped and the inner and outer toes are angled to the sides. The left front track (in the lower section a little below and to the left of the right front track) shows the four clawed toes, the C-shaped grouping of middle pads, and the two heel pads.

Mud is great, but winter is also fine for seeing squirrel family connections. In the photo below of red squirrel tracks in snow (bounding toward the top, of course) you see the same characteristic features you saw in the mud tracks. As sometimes happens, the heel area of the right rear foot (at the upper right of the photo) registered as a flattened area behind the middle pads. (If you look back at the first photo of the gray squirrel prints you’ll notice that the heel area of the left rear foot also made a slight impression.) There’s a variation in the arrangement of the front tracks, with the right front well behind but the left front farther forward. This kind of foot placement is often seen in squirrels, but is less common than the more four-square pattern.

Flying squirrels possess gliding membranes (the patagium) which extend between the front and rear legs, and because of this the rear feet can’t pass as far ahead of the front feet as they do in red or gray squirrels. In the next photo you see a bounding pattern made by a southern flying squirrel (oriented toward the top) in which the front prints are situated between rather than behind the rear prints. In northern flying squirrel trails the front prints often lie ahead of the rear ones. Another special flying squirrel trait is the thick covering of fur on the undersides of the feet. Because of this flying squirrel prints rarely show the crisp detail found in the tracks of other members of the squirrel family. But even with these differences, flying squirrel tracks will remind you of the tracks of other squirrels.

In the next image you see a bounding pattern made by a woodchuck. If you didn’t realize that woodchucks belong to the squirrel family, the familiar features of their tracks should make that clear. Woodchucks are more likely to walk than bound, and when a woodchuck does bound it usually places its front feet in a staggered pattern rather than even with each other, as in the photo. Nevertheless, the overall arrangement and the track details are consistent with those of its relatives.

To complete the picture for small rodents in the Northeast we need to add a few creatures that don’t strictly belong in the squirrel family but leave distinctly squirrel-like prints. These include white-footed mice, meadow voles, and their allies. I include mouse and vole allies because each one represents a group of closely related species which are difficult to distinguish from tracks alone.

First, let’s look at tracks of the white-footed mouse, shown below in a bounding pattern heading toward the upper right. In spite of its smaller size, the animal made tracks that are uncannily similar to the tracks in the first three photos. If I didn’t tell you that an individual rear print is just half an inch across you’d be hard pressed to tell these tracks from squirrel tracks.

Vole tracks also show striking similarities to the tracks we’ve already discussed–but with a few important differences. In the next photo you see tracks made by a meadow vole bounding from bottom to top. The track sequence, starting at the bottom, is: right rear, right front, left rear, left front. This staggered arrangement is common in vole trails and differs from the more consistent four-square bounding patterns usually seen in white-footed mice and tree squirrels. Voles can leave more regular bounding patterns, but they often move at something between a bound and a lope and their track patterns tend to be more variable. The toe impressions in vole tracks also tend to be more finger-like than the toes of mice. In spite of these differences the tracks of voles will remind you of mouse and squirrel tracks.

This is all well and good, you may say, but if these creatures are so similar to each other, how can I tell them apart? I’ve mentioned a few variations that can be helpful, but often the most useful trait is size. There’s a neat size progression, and although there’s some overlap between adjacent species it’s usually possible to make an identification with a few measurements combined with other clues. There are two dimensions to consider: track width (more reliable than track length) and bounding trail width (measured perpendicular to the direction of travel across the widest part of a bounding pattern). I’ll focus on the big picture rather than giving an exhaustive account of the numbers–detailed measurements can be found in any good tracking guide. White-footed mice and the smaller voles (woodland voles, for example) are the tiniest of the lot, and meadow voles are slightly larger. Chipmunks come next, and southern flying squirrels are slightly larger than chipmunks. Northern flying squirrels outweigh their southern kin, and red squirrels are larger yet. Gray squirrels beat out red squirrels, and woodchucks complete the series. These differences in body size are reflected in differences in track and trail dimensions, so a few measurements are usually sufficient to clinch an ID. Even when the tracks you’re dealing with are in the overlap zone there are usually other clues that can point toward an identification. And when all else fails, it’s okay to say you just can’t be certain. If you treat each situation as a learning experience, you’ll find yourself stumped less and less often.

Sorting Out the Small Rodents

Rodents are considered one of the most successful groups of mammals, so it’s not surprising that the northeast hosts many different kinds. They range in size from the tiny woodland vole (weighing an ounce or less) to the beaver (50 pounds or more). The small ones dominate, both in abundance and in their potential to confuse. Most of these little creatures are active in winter, so it’s a perfect time to get a handle on their distinguishing features.

The photo below shows a set of prints made by a red squirrel bounding from left to right. The five-toed rear tracks are nearly even with each other and set widely, their three middle toes parallel and their inner and outer toes diverging. The four-toed front tracks are set more narrowly and staggered, and their four toes are slightly splayed. Behind the toe impressions, the middle pads of both front and rear feet (analogous to the bumps over the knuckle joints in your palm and the ball of your foot) appear as clear indentations. The heel pads of the front tracks (like the heel of your hand) show in both right and left front prints, and the heel area of the right rear track (analogous to the heel of your foot) is a smooth elongation behind the middle pads.

Here’s a chipmunk group of four, with the direction of travel this time toward the top. The left front and rear tracks are partly superimposed, but the similarity to the tracks in the first photo is plain to see. This is what I call the rodent foot plan, and once you absorb it you’ll recognize it in other small rodents, including squirrels, chipmunks, mice, and voles.

But there are some variations which–if available–can be important in pinpointing an identification. The photo below came from the bounding trail of a southern flying squirrel, an animal similar to a chipmunk in body size (although lighter in weight). Compare the middle pads in the right rear tracks (the farthest to the right in each photo): in the chipmunk they’re well separated and form a sharp curve. The middle pads of the flying squirrel are closer together and form a gentle crescent.

If your reaction to that is, ‘you’ve got to be kidding!’ you’re not far off base. It’s a real difference, but snow conditions are rarely perfect enough to see that kind of detail. So how often can we be sure which small rodent made the tracks we’re seeing? Quite often, it turns out, because we have two additional diagnostic tools: trail width and habitual movement patterns. The tracks in the photo below, a white-footed mouse bounding toward the upper right and a gray squirrel bounding toward the lower right, are similar arrangements but are vastly different in size. In this case it’s easy to know which is which, but for less obvious differences, such as red squirrel versus gray squirrel, measurement of the overall width of the pattern, known as the trail width, can really help.

To measure the trail width of a bounding animal, imagine or mark lines parallel to the direction of travel which touch the outermost parts of the two rear tracks. Below you’ll see the same photo with lines delimiting the trail widths. Next, measure the distance between the two lines. The nice thing about this is that the trail widths of our most common small rodents fall into a simple size progression. In inches, trail widths for white-footed and deer mice measure 1 1/4-1 3/4; chipmunks, 1/1/2-2 3/4; red squirrels, 3-4 1/2; and gray squirrels, 4-6. At 1 3/4-3 inches the trail width for southern flying squirrels is similar to that of chipmunks, and northern flying squirrels, at 2 3/4-4 1/4 inches, overlap on the low side with red squirrels. Although trail width can be determined for any gait, the bounding gait so common in small rodents is especially suited to this measurement.

Habitual movement patterns are another useful tool for identifying small rodents. In the next photo a gray squirrel bounded at a good clip from bottom to top, leaving groups of four prints separated by relatively long distances. In each group of four the landing tracks of the smaller front feet are behind the take-off tracks of the larger rear feet. Bounding trails of red squirrels and chipmunks are similar in overall proportions. It’s not that these animals always make long leaps. If they’re moving slowly the distances between the groups of four can be smaller, and the hind feet may not pass as far ahead of the front feet. Compare the arrangement of the gray squirrel prints in the previous photo with that of the slower moving red squirrel in the opening illustration. The point is that the habitual travelling movement of these animals creates trails with characteristic four-track groupings and relatively large spaces between groups.

Compare the pattern above to the next photo, the trail of a southern flying squirrel, bounding from lower right to upper left. In this trail the larger rear prints are behind the smaller front ones, and the distance between the groups of four is smaller. In the trails of northern flying squirrels the rear tracks are often ahead of the front, but both species of flying squirrels have sacrificed strength for lightness and aerodynamic design and are unable to match the long leaps of their non-gliding relatives.

Snow depth can affect the foot placement of bounding rodents. To the white-footed mouse that made the tracks in the photo below the snow was fairly deep, so the groups of four are reduced to sets of two, each of the paired impressions made by sequential impacts of front and rear feet from the same side. All of the rodents I’ve been discussing do this when deep snow makes it more energy efficient. But even in these reduced patterns trail width can still be measured, as long as we make sure we’re looking at the actual tracks and not the larger openings around them. And like squirrels and chipmunks, mice make shorter leaps when moving less energetically. An example of mouse trails with consistently shorter leaps can be seen in the opening photo of last month’s article.

Meadow voles are chunkier and have shorter legs than white-footed mice, so they can’t make long leaps, but their trails are roughly as wide as those of mice. It’s not always easy to tell whether a bounding trail with short leaps was made by a vole or a mouse, but if the trail goes on long enough differences usually show up. A vole’s foot placement is rarely as even and foursquare as that of a mouse, and voles tend to make frequent shifts in gaits. It’s not unusual for an individual vole trail to vary among lopes, bounds, trots, overstep walks, and scurrying gaits that are difficult to categorize. In the next photo there’s a partly roofed vole tunnel meandering between the lower right and the top center. A vole traveled from the left side of the frame toward the tunnel in a bounding gait, with typical short leaps and uneven foot placement. The thin line in the center of the trail was made by the tail.

If you’ve made it this far in this treatise, you may feel like your brain is reeling. Believe it or not, I had to leave out many details, and I haven’t even addressed the issue of distinguishing small rodents from other small mammals. The important thing is to get started. Every time you work through a small rodent puzzle you’ll learn more. So be patient and persistent, and enjoy the eureka! moments when a few puzzle pieces fit together to form part of the larger picture.