The Behaviour of Ungulates and its Relation to Management: The Papers of an International Symposium held at the University of Calgary, Alberta, Canada, 2-5 November 1971. Volume 1

1. What they did

In November 1971, Valerius Geist and Fritz Walther brought together field workers who studied hoofed mammals (ungulates) and people who manage them in parks, reserves and game ranches, for a symposium at the University of Calgary, Canada. Their aim was to produce a reference book useful to both researchers and managers. Authors could revise their papers after the meeting, the papers were refereed, and some authors who could not attend sent papers too. The International Union for Conservation of Nature and Natural Resources (IUCN) published the result in 1974 in two volumes.

This entry covers Volume 1, which holds 25 papers. The editors describe it as dealing mainly with first attempts to bring together social behaviour and ecology, with theory and classification problems, with overviews of animal groups, and with new work on species not studied before. The papers include:

• Reviews and theory: mother and young relationships (Lent), expressive displays in fighting and courtship (Walther), social behaviour of the horse family (Klingel) and the pig family (Frädrich), social organization of 70 African bovids (Estes), and a theory linking ecology and behaviour in ungulate evolution (Geist).
• Field studies of single species: lesser kudu, Punjab urial, Walia and Alpine ibex, white rhinoceros, pronghorn (four papers), American bison, Newfoundland caribou, white-tailed deer (two papers), feral Soay sheep, vicuna, Coke’s hartebeest and collared peccary.
• Scent and experiments: scent glands in black-tailed deer, scent marking in captive Maxwell’s duikers, and the effect of foster rearing on fallow deer.

Volume 2, published at the same time, turns to ecology and management.

2. What they found

Each paper has its own findings. A few examples from Volume 1:

Mothers and young

• Most ungulates are either “hiders” or “followers”. Hiders, such as most deer, gazelles and many antelopes, leave their young lying hidden for long periods. Followers, such as horses, most large cattle relatives, sheep, muskox and caribou, stay close to their young from the start (Lent).
• Pronghorn fawns lie hidden up to 90% of the daytime for about the first three weeks. They choose their own beds, and seem to seek cover like the cover they were born in (Fichter).

Territories and marks

• About half of the African bovids (34 species) are gregarious and territorial; only about 10 seem not to be territorial at all (Estes).
• White rhinoceros bulls hold territories of about 2 km² for several years. Each territory has 20 to 30 dungheaps, where territorial bulls kick and scatter their dung; their spray-urination is most frequent along borders (Owen-Smith).
• Coke’s hartebeest males marked territories mainly with dung piles, often of several thousand pellets, most common along boundaries (Gosling).
• Vicuna families defend a feeding territory and a separate sleeping territory, with about five dung piles per hectare in feeding territories and seven in sleeping territories (Franklin).
• White-tailed deer bucks in Georgia made rubs and scrapes during the rut. Rubs seemed to express dominance; scrapes seemed to help bucks and does find each other. Bucks that found a doe’s scent trail at a scrape followed it with their nose near the ground and always found her quickly (Moore and Marchinton).

Movement

• Pronghorn use a wide range of gaits. When pressed they can jump barriers of up to eight feet, but they usually crawl under a fence if the bottom wire is at least 17 inches from the ground (Bullock).

3. What they concluded

• The editors see the papers as a reflection of the knowledge and the gaps of the time, not as final answers.
• They name weaknesses: the volumes lean heavily towards bovids, have little on deer other than caribou, and have no paper on how ungulates avoid predators.
• More research is needed on the basic biology of species not yet studied in the field (especially small tropical antelopes and deer, Asian and American deer, and rare forest and mountain species), and on animals in undisturbed ecosystems, so that natural forces rather than human management can be understood.
• Scientists and managers need to understand each other’s concerns, for the sake of conservation.

4. How tracking was or was not used

Most chapters rest on direct observation of animals. Tracks and signs feature strongly in a few:

• White-tailed deer (paper 22, pp. 447-456): the authors spent about 350 hours locating and studying field signs (rubs and scrapes), and surveyed deer sign in other habitats in Georgia and nearby states. At scrapes they usually found the tracks of the buck that made them, and later often tracks of smaller deer. Rubs stayed visible for at least several months. Trained hounds detected deer scent on rubbed trees several days later, and on branches over scrapes for at least four days.
• Dung and urine marks (papers 5, 7, 8, 15, 18, 24, 25): dungheaps, dung piles, middens, defecation stations, scrapes with paw marks, and urine-spray sites are described as territory or communication marks, with counts, densities and placement.
• Footprints and gaits (papers 3 and 12): black-tailed deer were seen sniffing the deep footprints of a fleeing deer; the pronghorn gait study describes where the hind foot lands relative to the front foot at slow and fast walks.
• Animals following scent: a caribou stag scented the ground where a herd had been the day before and followed the herd’s scent for a mile until he reached them (paper 20).

No trailing by the researchers is reported.

5. Tracker or observer certification

Not reported. No chapter describes the qualifications or testing of the people who found and interpreted tracks and signs; in paper 22 this was the authors themselves.

6. Other technologies, skills or innovations

• Radio-tracking of deer and rhinoceros, combined with deer conditioned in pens to tolerate a close observer before release.
• Trained hounds used to test whether rubs and marked branches carried deer scent.
• Frame-by-frame analysis of slow-motion film (80 frames per second) to study pronghorn gaits.
• Individual recognition from natural stripe patterns (lesser kudu) and photographic record cards (226 adult male and 37 adult female hartebeest).
• Experiments with captive animals: adding unfamiliar duikers to groups, and testing deer responses to whole and separated scent fractions.

7. Implications for conservation (from the paper)

• The editors call for short cuts to the conservation of endangered species, and for studies that reveal the principles of ungulate biology to help find them.
• The Walia ibex, found only in the Semien Mountains of Ethiopia, numbered about 150 animals in 1968, making it one of the most endangered mammals in the world (Nievergelt).
• For white rhinoceros, removing animals from certain “vacuum” zones is recommended, to control population growth without major social disruption (Owen-Smith).
• Herbicide spraying of sagebrush, the cover where pronghorn does prefer to give birth, raises the question of what happens to fawn survival where that cover is lost (Fichter).
• Fences with a bottom wire at least 17 inches above the ground let pronghorn crawl under them (Bullock).