# Animal Activity in Uncontrolled Terrestrial Communities as Determined by a Sand Transect Technique

> What if you could read every footprint left by a whole community, from beetles to bears, every two hours for a whole season? In 1960s Quebec, J. Roger Bider did just that with strips of fine sand, recording about 182,000 crossings and showing how mating, migration, rain and cold nights shape when and where animals move.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/1968-bider-animal-activity-uncontrolled-terrestrial/

## Citation and access

- **Authors:** J. Roger Bider
- **Year published:** 1968
- **Journal or source:** Ecological Monographs
- **Full citation:** Bider, J. R. 1968. Animal activity in uncontrolled terrestrial communities as determined by a sand transect technique. Ecological Monographs 38(4): 269-308.
- **DOI:** 10.2307/1948530
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.2307/1948530

## Tags

- **Species:** Mammals > American black bear; Amphibians > American bullfrog; Birds > American crow; Mammals > American mink; Mammals > American red squirrel; Birds > American robin; Amphibians > American toad; Birds > American woodcock; Birds > Black-capped chickadee; Birds > Blue jay; Mammals > Bobcat; Mammals > Brown rat; Mammals > Canada lynx; Birds > Chipping sparrow; Reptiles > Common garter snake; Mammals > Deer mouse; Mammals > Domestic cat; Mammals > Eastern chipmunk; Mammals > Eastern grey squirrel; Birds > European starling; Amphibians > Green frog; Amphibians > Grey tree frog; Invertebrates; Amphibians > Jefferson salamander; Mammals > Meadow jumping mouse; Mammals > Meadow vole; Mammals > Moose; Mammals > Mountain lion; Mammals > Muskrat; Mammals > North American porcupine; Mammals > North American river otter; Birds > Northern flicker; Mammals > Northern flying squirrel; Amphibians > Northern leopard frog; Mammals > Northern short-tailed shrew; Amphibians > Northern two-lined salamander; Birds > Ovenbird; Amphibians > Pickerel frog; Mammals > Raccoon; Mammals > Red fox; Reptiles > Red-bellied snake; Reptiles > Ring-necked snake; Birds > Ruffed grouse; Birds > Savannah sparrow; Reptiles > Smooth green snake; Mammals > Snowshoe hare; Birds > Song sparrow; Mammals > Southern red-backed vole; Amphibians > Spring peeper; Mammals > Star-nosed mole; Mammals > Stoat; Mammals > Striped skunk; Birds > Veery; Mammals > White-tailed deer; Birds > White-throated sparrow; Amphibians > Wood frog; Mammals > Woodchuck; Mammals > Woodland jumping mouse
- **Region:** North America > Canada
- **Tracking:** Gaits; Track casts; Track counts; Tracking; Tracks; Tracks and signs identification
- **Behaviour and ecology:** Behaviour; Dispersal; Feeding; Food caching; Habitat selection; Hunting; Migration; Movement
- **Methods and technology:** Analysis of variance (ANOVA); Chi-square test; Descriptive statistics; Direct observation
- **Data type:** Modelling or theoretical; New field data
- **Observer qualification:** Not reported
- **Discipline:** Behaviour and ecology; Wildlife tracking
- **Year published:** 1968

## Detailed keywords

- **Species:** Studied: the whole ground-dwelling animal community crossing the transects, recorded as invertebrates (arthropods 1/8 inch long and larger, earthworms, snails, roundworms and horsehair worms) and 58 vertebrate species grouped into 38 categories. Mammals include porcupine (Erethizon dorsatum), white-tailed deer (Odocoileus virginianus), moose (Alces alces), snowshoe hare (Lepus americanus), northern flying squirrel (Glaucomys sabrinus), grey squirrel (Sciurus carolinensis), red squirrel (Tamiasciurus hudsonicus), eastern chipmunk (Tamias striatus), deer mouse (Peromyscus maniculatus), meadow and woodland jumping mice (Zapus hudsonius, Napaeozapus insignis), red-backed vole (Clethrionomys gapperi), meadow vole (Microtus pennsylvanicus), woodchuck (Marmota monax), brown rat (Rattus norvegicus), muskrat (Ondatra zibethicus), shrews (Sorex sp., Blarina brevicauda), star-nosed mole (Condylura cristata), striped skunk (Mephitis mephitis), raccoon (Procyon lotor), Canada lynx (Lynx canadensis), bobcat (Lynx rufus, from 1965), red fox (Vulpes fulva), ermine (Mustela erminea), mink (Mustela vison), river otter (Lutra canadensis), domestic cat (Felis domestica), mountain lion (Felis concolor, a record for the region) and black bear (Ursus americanus). Birds include ruffed grouse (Bonasa umbellus), American woodcock (Philohela minor), northern flicker (Colaptes auratus), American crow (Corvus brachyrhynchos), blue jay, American robin, starling, veery, thrushes, ovenbird, cowbird, black-capped chickadee and chipping, white-throated, savannah and song sparrows. Amphibians: American toad (Bufo americanus), green, leopard, wood, pickerel and bull frogs (Rana spp.), spring peeper and grey tree frog (Hyla spp.), two-lined salamander (Eurycea bislineata) and Jefferson salamander (Ambystoma jeffersonianum). Reptiles: red-bellied snake (Storeria occipitomaculata), garter snake (Thamnophis sirtalis), ring-necked snake (Diadophis punctatus) and green snake (Opheodrys vernalis). Present on the site but never seen crossing: red-backed salamander (Plethodon cinereus). Said to be trapped in similar habitats but not recorded: American marten (Martes americana), fisher (Martes pennanti), long-tailed weasel (Mustela frenata) and wolf (Canis lupus).
- **Location and habitat:** About 2 miles north of the village of Lac Carré, Terrebonne County, Quebec, Canada (latitude 46° 09’, longitude 74° 29’), in the mid-Laurentians about 65 miles northwest of Montreal, at 1000 to 1200 ft. Old fields, a forest edge and mixed forest of maple, yellow birch, beech and softwoods (spruce and balsam fir), on post-glacial terraces beside a small creek, with a kettlehole and frost pocket along the ecotone transect. Humid continental climate with cool summers.
- **Topics:** sand transects (track beds) as a community-level method; animal activity rates; spatial use of woods, fields and forest edge; edge effect and an “activity vacuum” behind the forest edge; runways; homesites and feeding paths; daily (diel) activity peaks; seasonal phenology of activity (maintenance, mating, storage, migration, dispersal); hyperactivity; effects of weather systems, rain, cloud cover and temperature thresholds; interactions between species (“biofriction”); track identification criteria for voles and squirrels; plaster casts of sand prints as a reference collection; transect canopy for tracking in rain.
- **Statistics or models:** Repeated one-way analysis of variance (ANOVA) to group sections and periods into “activity levels”; chi-square test; percentages of daily activity per hour; sign (plus and minus) comparisons of day-to-day changes between species; a theoretical model of how much activity should rise as litters disperse (Fig. 18); a synoptic outline of a four-level hierarchy of factors affecting activity (Fig. 23). Inconsistency: the text says 16 of 20 species used the deeper woods at random, but the woodland summary on the same page says random use by “18-20 species”.
- **Data type:** New field data: three transects tracked from 1962 to 1964. In 1963, four discrete tracking periods (late June, late July, late August to early September, mid-October). In 1964, continuous tracking under a canopy for 165 days (every 2 hours from 1 June to 10 September, then at least twice a day to 15 November), with weather records. About 182,000 crossings and 20,000 climate readings. Some remarks draw on later data (1965 to 1967). Includes a theoretical model.

## Synopsis

### 1. What they did

Bider wanted a single, simple method that could record the activity of a whole community of ground-dwelling animals, from insects and worms to mammals and birds, in natural conditions. Every animal moving on the ground leaves a trail in fine sand, so he built strips of sand about 3 ft wide and several hundred feet long, and noted the location and direction of every track crossing them at regular intervals.

- The transects. Near Lac Carré, Quebec, he built three: an ecotone transect (1962) running 450 ft through an open field and 300 ft into the woods, a 783-ft field transect and a 450-ft woods transect (both 1963). Soil was cleared of roots and pebbles, compacted and topped with fine sand. After each reading the sand was swept smooth with a soft push broom.

- Reading the tracks. Tracks were read every 2 hours, day and night (at night by gas lantern), and recorded in 10-ft units on data sheets.

- 1963: four tracking periods. Late June, late July, late August to early September, and mid-October, with each transect divided into 150-ft sections to compare where animals were active.

- 1964: continuous tracking. A polyethylene canopy, 6 ft wide and suspended 6 ft high, kept rain off the ecotone transect so it could be tracked continuously for 165 days, alongside detailed weather records.

He did not try to count individual animals: the measure was the number of crossings, which reflects activity rather than population size.

### 2. What they found

The community

- The study produced about 182,000 crossings. The community comprised invertebrates and 58 vertebrate species, grouped into 38 categories where species were hard to tell apart.

- The ecotone transect recorded about twice as many species as the field or woods transects (51 against 32 and 30); with the canopy and continuous tracking, 59 species and 9 more were recorded.

- A 150-ft section recorded as many species as a whole 783-ft transect in similar habitat. Both lynx species and a mountain lion crossed, the latter a record for the region.

Use of space

- In the woods, most species eventually used the whole area at random (16 of 20 species used the deeper sections equally). Activity was often higher in the first 150 ft inside the forest edge and lower in the next 150 ft, an “edge and vacuum” effect.

- In the fields, activity was more patchy: many species used pockets of the field, and activity was lowest near the field edge.

- Deer mice moved along set paths from a homesite to feeding areas. Meadow voles and shrews crossed mostly at about 10 major runways (82% and 83% of their crossings), which took up less than 4% of the field transect.

- Snowshoe hares shifted during the summer from the deep woods toward the forest edge and into the fields, following the food supply. Garter snakes seemed to move down to the fields to give birth and back to the woods to hibernate.

- Red squirrels and chipmunks used separate areas when both were most active in late summer, the squirrels carrying cones from a spruce and a chipmunk carrying leaves to a hole.

Time of activity

- Peaks of activity of different species were most spread out on warm July nights, and squeezed into the first hours after dusk on cold nights. In autumn, arhythmic animals (voles, shrews, snakes) shifted their activity from night to day.

- Through the season, activity rose and fell with life events: short bursts during mating (chipmunks in a “frenzy” along the forest edge; five hares chasing in daylight), high steady activity while storing food, and sudden surges during migrations of frogs, salamanders, snakes and birds.

- Weather shaped activity within each season. Activity was lowest the first day after a front; diurnal animals peaked on the last clear day and nocturnal animals on warm cloudy nights. Rain suppressed insect activity, stimulated meadow voles, and did not change shrew activity. Toads were hardly active when dusk temperatures were not above 50°F.

- Ermine crossing many times in one day left mostly a crouched gait instead of their normal hopping gait.

### 3. What they concluded

- Activity in a natural community is controlled at four levels, each setting limits for the next: the year (population size), the season and its life events, each weather system, and individual climatic events.

- Simpler animals respond mainly to physical conditions such as temperature and rain; more advanced animals depend more on behaviour. Bider called this a “physio-ethological activity gradient”.

- Activity data from a single day are worthless. Analyses should compare similar days within one year, within one seasonal event and from similar points in weather systems.

- One full weather system is the minimum time needed to detect the basic species of a community in a given season.

- Bider considered the sand transect his greatest contribution: in his view, the most practical, simple, economic and least biased way to get copious data on animal activity.

### 4. How tracking was or was not used

Tracking was the method. Every track crossing the sand was identified and recorded by location and direction. Identification used Murie’s Field Guide to Animal Tracks; the author notes that Murie’s drawing for red-backed voles is wrong (the drawings were not Murie’s own). Red-backed and meadow voles were separated by print size and depth, and by gait (meadow voles walk; red-backed voles use a triangular hopping gait), with decisions on new runways postponed until the animal was seen; the author estimates this kept errors under 1%. Chipmunks, red squirrels, grey squirrels and woodchucks were separated by stride width, print weight and distance between jumps. Shrews and jumping mice could not be told apart reliably and were grouped by family. Questionable tracks were circled and timed in the sand for checking. A reference collection of real prints was made by spraying the sand with lacquer and setting it on plaster of Paris.

### 5. Tracker or observer certification

Not reported. The author says that identifying tracks “is a question of practice and experience”. Student assistants did much of the reading; no training, certification or test of their skill is described. The author notes that some squirrel tracks gave the assistants the most trouble.

### 6. Other technologies, skills or innovations

- A transect canopy of polyethylene film that allowed continuous tracking through rain and limited dew.

- A sand recipe and maintenance routine: fine sand (particles of about 0.1 mm) to record small details, raking to dry the track after rain, and herbicide to control weeds.

- Weather recording with a time-lapse movie camera (1962 to 1963), then a rain and snow gauge, dew gauges, a wind instrument and a pyrheliometer.

- Lacquer and plaster casts of sand prints for teaching and checking.

### 7. Implications for conservation (from the paper)

- The edge and vacuum pattern at the forest edge may matter in applied ecology, where strip cutting is used to produce a greater abundance of animals.

- The yearly number of crossings by transient animals during the dispersal phase might be a good index of a population’s success in rearing young.

### 8. Implications for tracking (Original Wisdom's observations)

_These are Original Wisdom's observations, not the authors'._

- This early study shows how much a prepared tracking surface can reveal: when, where and under what weather a whole community moves, including insects, worms, frogs and snakes that are rarely studied by tracks.

- The author’s vole and squirrel criteria combine print size, depth, gait and pattern, and he postponed doubtful identifications until he could see the animal. Both remain useful habits.

- Observer skill was not tested, so the accuracy of identifications for the harder groups is unknown.

## Cite this summary

Lawrence, K. 2026. Summary of: Animal Activity in Uncontrolled Terrestrial Communities as Determined by a Sand Transect Technique (Bider, 1968). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/1968-bider-animal-activity-uncontrolled-terrestrial/