# 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

> In 1971, ungulate researchers from around the world met in Calgary to share what they knew about how hoofed animals live together. The first volume of their papers covers everything from hiding fawns and fighting displays to rhino dungheaps, hartebeest dung piles and the rubs and scrapes of white-tailed deer. It is a rich, early record of the signs ungulates leave for each other, and for trackers.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/1974-geist-walther-behaviour-ungulates-relation-management/

## Citation and access

- **Authors:** Valerius Geist and Fritz R. Walther
- **Year published:** 1974 (symposium held in 1971)
- **Journal or source:** IUCN Publications new series No. 24, Volume 1. International Union for Conservation of Nature and Natural Resources (IUCN), Morges, Switzerland. Book, 511 pages.
- **Full citation:** Geist, V. and Walther, F. (eds.) 1974. 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. IUCN Publications new series No. 24. Morges, Switzerland: International Union for Conservation of Nature and Natural Resources. 511 pp. ISBN 2-88032-013-5.
- **Access status:** Hosted (freely available elsewhere)
- **PDF:** https://www.originalwisdom.com/wp-content/uploads/2026/11/1974_Geist_and_Walther_the-behaviour-of-ungulates-and-its-relation-to-management-volume-1.pdf

## Tags

- **Species:** Mammals > African wild ass; Mammals > Alpine ibex; Mammals > American bison; Mammals > Asiatic wild ass; Mammals > Black-tailed deer; Mammals > Caribou; Mammals > Coke’s hartebeest; Mammals > Collared peccary; Mammals > Fallow deer; Mammals > Grant’s gazelle; Mammals > Grevy’s zebra; Mammals > Lesser kudu; Mammals > Maxwell’s duiker; Mammals > Mountain zebra; Mammals > Plains zebra; Mammals > Pronghorn; Mammals > Przewalski’s horse; Mammals > Soay sheep; Mammals > Thomson’s gazelle; Mammals > Urial; Mammals > Vicuña; Mammals > Walia ibex; Mammals > Warthog; Mammals > White rhinoceros; Mammals > White-tailed deer; Mammals > Wild boar
- **Region:** North America > Canada; Africa > Ethiopia; Africa > Kenya; Asia > Pakistan; South America > Peru; Africa > South Africa; Europe > Switzerland; Europe > United Kingdom; North America > United States
- **Tracking:** Beds; Gaits; Scat; Sign surveys; Tracking; Tracks; Tracks and signs identification; Urine
- **Behaviour and ecology:** Behaviour; Communication; Home range; Movement
- **Methods and technology:** Chi-square test; Descriptive statistics; Direct observation; Literature review; Radio telemetry
- **Data type:** Modelling or theoretical; New field data; Review
- **Observer qualification:** Not reported
- **Discipline:** Behaviour and ecology; Wildlife tracking
- **Year published:** 1974

## Detailed keywords

- **Species:** Studied (hoofed mammals, the ungulates, wild, captive, feral and domestic): many species across the volume, including white-tailed deer (Odocoileus virginianus), black-tailed deer (Odocoileus hemionus columbianus), fallow deer (Dama dama), caribou (Rangifer tarandus), pronghorn (Antilocapra americana), American bison (Bison bison), Punjab urial (Ovis orientalis punjabiensis), feral Soay sheep, Walia ibex (Capra walie), Alpine ibex (Capra ibex), Maxwell’s duiker (Cephalophus maxwellii), lesser kudu (Tragelaphus imberbis), Coke’s hartebeest (Alcelaphus buselaphus cokei), Thomson’s gazelle and Grant’s gazelle, white rhinoceros (Ceratotherium simum), vicuna (Vicugna vicugna), collared peccary (Dicotyles tajacu), wild boar (Sus scrofa), warthog (Phacochoerus aethiopicus), plains zebra (Equus quagga), mountain zebra (Equus zebra), Grévy’s zebra (Equus grevyi), wild horse (Equus przewalskii), African wild ass (Equus africanus) and Asiatic wild ass (Equus hemionus). One review chapter (Estes) surveys the social organization of 70 African bovids (69 antelopes and the Cape buffalo); another (Geist) covers the evolution of ungulates from the fossil record. Predators named in the hartebeest chapter: lion and cheetah. Goats were used as foster mothers for fallow deer fawns, and trained hounds tested deer scent marks.
- **Location and habitat:** Field studies in this volume come from: Georgia, United States (Piedmont pine and pine-hardwood forest near Athens, and Jones Island in the Okefenokee National Wildlife Refuge swamp); National Bison Range, Montana, and Wind Cave National Park, South Dakota, United States; Catalina Island, California, United States; Rob and Bessie Welder Wildlife Refuge, Texas, United States; Upper Pahsimeroi drainage, central Idaho, United States (sagebrush); Arizona, United States (Pinal Mountains); southern Alberta grasslands, Canada; Newfoundland, Canada, and Mt. Albert, Gaspé Peninsula, Quebec, Canada; Umfolozi-Corridor-Hluhluwe game reserve complex, Zululand, South Africa; Tsavo National Park (East), Kenya; Nairobi National Park, Kenya (short grass plains, Acacia drepanolobium scrubland and valley slopes); Semien Mountains, Ethiopia; Salt Range, Pakistan (a private sanctuary); Pampa Galeras National Vicuna Reserve, southern Peru; Hirta, St. Kilda, Scotland, United Kingdom; Swiss Alpine ibex colonies, Switzerland; Bronx Zoo, New York, United States (captive duikers). Symposium venue: University of Calgary, Alberta, Canada.
- **Topics:** Edited symposium volume (IUCN). Volume 1 papers, with book page ranges: (1) Lent, mother-infant relationships, hiders and followers, pp. 14-55; (2) Walther, expressive behaviour in combat and courtship of horned ungulates, pp. 56-106; (3) Müller-Schwarze, social functions of scent glands in black-tailed deer and pronghorn, pp. 107-113; (4) Ralls, scent marking in captive Maxwell’s duikers, pp. 114-123; (5) Klingel, social behaviour of the Equidae, pp. 124-132; (6) Frädrich, behaviour in the Suidae (pigs), pp. 133-143; (7) Sowls, social behaviour of the collared peccary, pp. 144-165; (8) Estes, social organization of the African Bovidae, pp. 166-205; (9) Leuthold, home range and social organization of lesser kudu, pp. 206-234; (10) Geist, ecology and behaviour in the evolution of ungulates (dispersal theory), pp. 235-246; (11) Gilbert, foster rearing and adult social behaviour in fallow deer, pp. 247-273; (12) Bullock, locomotion and gaits of pronghorn, pp. 274-305; (13) Schaller and Mirza, behaviour of Punjab urial, pp. 306-323; (14) Nievergelt, rutting and grouping in Ethiopian (Walia) and Alpine ibex, pp. 324-340; (15) Owen-Smith, social system of the white rhinoceros, pp. 341-351; (16) Fichter, bedding behaviour of pronghorn fawns, pp. 352-355; (17) Bromley and Kitchen, courtship in the pronghorn, pp. 356-364; (18) Kitchen and Bromley, agonistic behaviour of territorial pronghorn bucks, pp. 365-381; (19) Lott, sexual and aggressive behaviour of American bison, pp. 382-394; (20) Bergerud, rutting behaviour of Newfoundland caribou, pp. 395-435; (21) Brown, social organization in male groups of white-tailed deer, pp. 436-446; (22) Moore and Marchinton, marking behaviour and its social function in white-tailed deer, pp. 447-456; (23) Grubb, mating activity of rams in feral Soay sheep, pp. 457-476; (24) Franklin, social behaviour of the vicuna, pp. 477-487; (25) Gosling, social behaviour of Coke’s hartebeest, pp. 488-511. Themes: social organization, territoriality, dominance, rut and courtship, mother-infant bonds, imprinting, expressive displays, scent glands and scent marking, signposts, home range, grouping, evolution, locomotion and gaits, management of wild and captive ungulates, endangered species. Chapters with tracks and signs content: antler rubs, scrapes, marked branches, tracks at scrapes and scent trails of white-tailed deer (22); white rhinoceros dungheaps with scattered dung and urine-spray sites concentrated along territory borders (15); hartebeest boundary dung piles of several thousand pellets (25); vicuna dung piles, their density and weight in feeding and sleeping territories (24); pronghorn buck scrapes with paw marks, feces and urine, and thrashed vegetation (18); deer sniffing fecal pellets, urine and footprints, and forehead-rub marks (3); footfall placement of fore and hind feet at different gaits, and fence crossing (12); fawn beds and bedding cover (16); caribou bush thrashing and a stag following a herd’s scent trail on the ground (20); bison wallows (19); urial horning of shrubs (13); dung middens and preorbital gland marks of small antelopes (8); zebra and wild ass dung piles up to 40 cm high (5); peccary defecation stations and droppings along trails (7). Note on the paper: the contents page lists the co-author of paper 22 as “L. Marchinton”, while the paper itself gives “R. Larry Marchinton”.
- **Statistics or models:** Varies by chapter. Mostly descriptive statistics (counts, percentages, means, group sizes, home range areas). Chi-square tests (for example on Thomson’s gazelle threat encounters, paper 2); Spearman rank correlation, Wilcoxon and chi-square tests (ibex, paper 14); analysis of variance and t tests (duikers, paper 4); t tests and rank correlations (fallow deer, paper 11); life tables (Soay sheep, paper 23); frame-by-frame analysis of slow-motion film (pronghorn gaits, paper 12); a theoretical model of ungulate evolution, the dispersal theory (paper 10).
- **Data type:** New field data, reviews and theory. Examples of data collection dates: white rhinoceros January to June 1966 and November 1968 to September 1971; vicuna May 1967 to June 1971; urial 6 October to 9 November 1970; white-tailed deer marking 1968 (Athens) and 1969 (Okefenokee); male white-tailed deer groups 1969 to 1970; bison breeding seasons 1966 to 1971; Newfoundland caribou rut in ten seasons between 1952 and 1964; pronghorn fawns over 16 years. Several chapters are reviews (papers 1, 2, 5, 6, 8) or theory (paper 10). This PDF holds Volume 1 only (papers 1 to 25, book pages 1 to 511); Volume 2 (papers 26 to 56, pages 512 to 940, management chapters) is a separate file. Book page 99 (in the appendix tables of paper 2) is not in the scan. Year: the symposium was in 1971 and the book was published in 1974; the old library listing used 1971.

## Synopsis

### 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).

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

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

- Much of what animals “say” to each other is written on the ground: rubs, scrapes, dung piles, middens, urine sites and wallows. Several chapters show that where these signs are placed (along borders, at trail junctions, under marked branches) carries meaning, which is useful for trackers reading territories and the rut.

- The white-tailed deer chapter is an early example of researchers combining radio-tracking, direct observation and tracks and signs to work out what a sign means.

- The animals themselves are trackers: deer, caribou and pronghorn sniff footprints, urine and scrapes, and follow scent trails to find each other.

## Cite this summary

Lawrence, K. 2026. Summary of: 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 (Geist and Walther, 1974). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/1974-geist-walther-behaviour-ungulates-relation-management/