Gundis

(Ctenodactylidae)

Class Mammalia

Order Rodentia

Suborder Sciurognathi

Family Ctenodactylidae

Thumbnail description
Small, thickset rodents with soft, thick fur, large, blunt heads, short, rounded ears, large eyes, short legs and short, furry tails.

Size
Body 5.5–9.5 in (14–24 cm); tail 0.4–2.4 in (1–6 cm); 6.0–10.2 oz (170–290 g)

Number of genera, species
4 genera; 5 species

Habitat
Rocky areas in deserts or semidesertic ranges.

Conservation status
Vulnerable: 1 species

Distribution
Northern Africa, from Mauritania and Morocco east to Eritrea and Somalia.

Evolution and systematics

The gundis (family Ctenodactylidae) are usually classified within the suborder Hystricomorpha but they may have affinities with the Paramyids from the Eocene and, as the sciurid rodents, may also be descended from the Paramyids. The gundis could perhaps be placed in a separate superfamily (Ctenodactyloidea) within the Sciuromorpha (where they are included here) rather than the Hystricomorpha.

The family contains five species in four genera: Pectinator, Felovia, Massoutiera, and Ctenodactylus.

Hartenberger, in 1985, concluded that the Ctenodactylidae might be one of the oldest rodent families, having diverged from other rodent taxa in the early Eocene. Molecular data analysis of amino acid myoglobin sequences performed by Beintema et al. in 1991 provided supporting evidence for the hsytricognathous rodents and the Ctenodactylidae being early offshoots of the order Rodentia, although it proved impossible to decide whether they share a common ancestor.

The earliest fossils assigned to the Ctenodactylidae date from the middle Eocene of Asia. There are also Oligocene remains from central Asia and Miocene and Pleistocene remains from Asia, Sardinia, Sicily, and North Africa. The four living genera originated in Africa, where they are known only from the Recent period.

Physical characteristics

Gundis are small, stocky rodents with short, furry tails and short legs. They resemble guinea pigs (Cavia) in external appearance. Females are on average larger than males.

The head is large and blunt, the eyes large and the vibrissae long. The ears are short and rounded, and in some species have a protective fringe of hairs around their inner margin. In the mzab gundi (Massoutiera mzabi) the ears are flattened against the head and do not move.

The hind feet are longer than the forefeet. Each foot has four digits and on the hind feet the two inner digits have stiff bristles that form a comb: the family name translates as "comb-fingered." All digits have small, sharp claws. The two Ctenodactylus species have very small wispy tails but the other three species have longer, fan-like tails.

The fur is soft, silky, and dense. In Speke's pectinator (Pectinator spekei) the skin is thin and easily torn. Crouched on a rock with the wind blowing through their fur, gundis look like powder puffs. The cuticular scales on the hairs are unusual, being narrow and petal-like.

Gundis vary in color from grayish or buff to chestnut or yellowish red, with paler (usually whitish) underparts. The color of each species matches the rocks in which it occurs.

The skull is flattened, with broad frontals and relatively well-developed supraorbital ridges. The jugals have horizontal and vertical branches, and the vertical reaches the enlarged lacrimal. The bullae and mastoids are inflated, and the skull appears to broaden posteriorly. The dental formula is i1/1, c0/0, pm1/1 or 2/2, m3/3 = 20 or 24. The cheek teeth are flat-surfaced and ever-growing. Gundis have a flexible ribcage that allows them to squeeze into crevices.

The family is unusual in that females have a cervical pair of mammae in addition to the pair placed laterally on the anterior thorax.

Distribution

Gundis are confined to northern Africa. Speke's pectinator occurs from Eritrea and Ethiopia east to Somalia, while the other four species occur further west, in the Sahara region. The mzab gundi is confined to the central Sahara from Algeria and Mali to Libya and Chad, while the felou gundi (Felovia vae) occurs in Mauritania and Mali. The two Ctenodactylus species have overlapping ranges, the desert gundi (C. vali) ranging from Morocco through Algeria to Libya, while the North African gundi (C. gundi) also extends into Tunisia.

Habitat

Gundis are found on rocky outcrops, screes, hills, cliffs, and mountains in desert, subdesert, or desert edge habitats between sea level and 8,200 ft (2,500 m). Geologically, the rocks may be of any age and of any type from recent lava flows to ancient folded sandstone. In some areas, road building sites have also been occupied. Rocks must not be too large, and fissures, crevices, and caves are essential for permanent and temporary shelters, while ledges, flat rocks, and boulder tops are used for sunbathing. Gundis often favor a site with an easterly aspect, exposed to the morning sun, and ideally some part of the site should also catch the evening sun.

Behavior

Gundis do not excavate burrows but live in natural rock crevices or caves. They are gregarious, living in colonies that vary in density from the mzab gundi's 0.12 per acre (0.3 per ha) to over 40 per acre (100 per ha) for Speke's pectinator. Density is related to food supply and the nature of the terrain. Colony size varies from 16,150–21,500 sq ft (1,500– 2,000 sq m) in Speke's pectinator to 26,900 sq ft (2,500 sq m) in the desert gundi.

Within colonies there are family territories occupied by a pair and their juveniles or by several females and offspring. Gundis do not make nests, and shelters are often temporary. Permanent shelters may be occupied for many years. Characteristically, a shelter retains the day's heat through the cold night and provides cool conditions in the heat of the day.

The large eyes might suggest nocturnal habits, but gundis are diurnal, moving rapidly from bright sunlight to deep shade in rock crevices. They often move slowly but can run quickly, with the belly almost touching the ground. They are shy and wary, and rely mainly on speed and agility to squeeze into crevices and holes to escape from predators. Rough friction pads on the soles of the feet assist in climbing on rocks, and gundis can ascent almost vertical surfaces, keeping the body pressed close to the rock face. All gundis thump with their hind feet when alarmed. Their flat ears allow them good all-round hearing and the hearing is acute.

When the weather is cold, wet, or windy, activity is restricted and the animals may not emerge at all. In winter, gundis pile on top of one another for warmth, with juveniles shielded by their mother or draped across the back of her neck. Gundis are not known to hibernate or estivate.

Gundis normally emerge at first light and remain active for up to five hours. Activity declines during the hot part of the day but increases again in the 2–4 hours before dusk.

The three species with fan-like tails use the tail for balance, while in Speke's pectinator the tail is also used in social displays.

Grooming is a common activity. The combs on the hind feet are used for grooming and scratching, and the rapid circular scratch of the rump with a hind foot is characteristic of gundis. The fur does not repel water and in wet weather it sticks together in tufts. The animals take particular care to ensure that the fur remains loose.

Vocalizations are varied, and each species has its own repertoire. Calls vary from the infrequent chirp of the mzab gundi to the relatively complex chirps, whistles, and chuckles of Speke's pectinator. In their habitat their low-pitched calls carry well. Short, sharp calls warn of predatory birds and cause all nearby gundis to take cover. Longer calls warn of ground predators and inform the predator it has been spotted. The felou gundi's harsh "chee" call continues as long as the predator remains in the vicinity.

Long complex chirps or whistles serve for recognition or greeting. The two Ctenodactylus species, whose ranges over-lap, have very different calls that aid in species recognition: the North African gundi chirps, the desert gundi whistles.

Gundis "play possum," exhibiting an immobile, trancelike state when threatened by a predator such as a snake, lizard, fox, jackal, or cat, most of which hunt by sight. The gundi lies flat and completely immobile, and may stop breathing for up to a minute. Gundis do not bite when handled.

For small desert mammals, gundis are unusually active in the daytime. In the early morning they stretch out flat on their stomachs to sunbathe until the temperature rises above 20°C (68°F), when they forage. After feeding, they again flatten themselves on the warm rocks to keep their bodies warm and to speed digestion: a way of making the best use of scarce food. When the temperature reaches 32°C (90°F) the gundis take shelter under rocks and do not emerge until the temperature drops in the afternoon. When long foraging expeditions are necessary, gundis alternate feeding in the sun with cooling off in the shade. In extreme drought, gundis eat at dawn when plants contain the most moisture.

Gundis have communal dunghills, some of which may have been in use for many years, and finding such a latrine may be the easiest way to ascertain the animals' presence in an area.

Feeding ecology and diet

These animals are entirely herbivorous. Their diet includes the leaves, stalks, flowers, and seeds of almost any desert plant, including grasses and acacia trees. Preferred plants include those in the families Cruciferae and Compositae. Gundis are not adept at gnawing and their incisors lack the hard orange enamel characteristic of most rodents.

Food is often scarce and gundis may forage over great distances, sometimes traveling 0.6 mi (1 km) in a morning. Regular foraging is essential because food is not stored and fat reserves are not accumulated in the body.

Most gundis do not drink but obtain all the water they need from plants; their kidneys have long tubules for absorbing water and under extreme conditions their urine can be concentrated, although this emergency response can only be sustained for a limited period.

Reproductive biology

Most species produce young between January–March and June, with the anestrus period extending for 6–8 months thereafter. The estrus cycle is 23–25 days and the gestation period 55–56 days. The litter size is 1–3 and most species apparently produce only one litter per year. The one exception to this seasonal cycle is Speke's pectinator, which appears to be a more opportunistic breeder, captive females being in anestrus only in July.

The young are born fully furred with their eyes open and are brought out into the open within a few hours of birth. They are left in a rock shelter while the mother forages and their continuous chirruping helps the mother relocate them in their temporary shelter. The female has been observed to carry small young in her mouth by the skin of their necks.

There are few opportunities to suckle, and young are fed chewed leaves from the start. They are weaned at about 4 weeks of age. Weaning probably starts early because the mother can produce little milk in the dry heat of the desert. Sexual maturity is attained at 8–12 months.

Longveity in the wild is estimated at 3–4 years; in captivity longevity of 10 years is recorded for Speke's pectinator.

Conservation status

The felou gundi is classed as Vulnerable because of a decline in its overall range and habitat. Other gundis are not globally threatened, although some may be locally threatened by hunting for food, disturbance from close human settlements, and predation by domestic cats and dogs.

Significance to humans

Gundis are well known to local people and "gundi" is their Arabic name, although it is also applied to other rodents of similar appearance. Twilight, in the common speech of Arabs, is known as "the hour when the gundi comes out." Gundis are hunted for food by some North African tribes. Although not regarded as pests, gundis could be potentially destructive to crops and gardens.

Species accounts

List of Species

Speke's pectinator

Pectinator spekei

taxonomy

Pectinator spekei Blyth, 1856, Somalia.

other common names

English: Bushy-tailed gundi; French: Pectinator de Speke; German: Buschschwanzgundi.

physical characteristics

Head and body length, 5.5–7.5 in (14–19 cm); tail 1.6–2.4 in (4–6 cm); and about 6.3 oz (178 g) for captive females. Gray, tinged brown or black; underparts grayish white; tail bushy.

distribution

Eritrea, eastern and southern Ethiopia, Djibouti, and northeastern Somalia.

habitat

Rocky cliffs and outcrops in desert or semidesert, often with hyrax Procavia; occurs from sea level to 5,900 ft (1,800 m).

behavior

Shelters in rock crevices. Diurnal, emerging to feed in early morning, peak activity 2–4 hours after dawn. It often basks in the sun. Vocalisations are a relatively complex range of chirps, chuckles and whistles; utters a whistling call on the approach of a predator; the normal call is described as a long, drawn-out "whee whee." Colonies of 16,150–21,500 ft² (1,500–2,000 m²) with well-defined boundaries have been recorded; may occur at densities of over 40 individuals/acre (100/ha). Bushy tail used in social displays.

feeding ecology and diet

Eats only plant material, including dry grass stalks and seeds, the leathery green leaves of Cadaba rotundifolia bushes, and the leaves of Acacia senegal and the long-spined A. seyal.

reproductive biology

Young born August–September. Captive females in anestrus only in July; possibly a more opportunistic breeder than Ctenodactylus and Massoutiera. Estrus cycle averages 22.7 days; litter size in captivity one, occasionally two. Life span in captivity 10 years.

conservation status

Not globally threatened.

significance to humans

None known.

Mzab gundi

Massoutiera mzabi

taxonomy

Ctenodactylus mzabi (Lataste, 1881), Ghardaia, northern Algeria.

other common names

English: Lataste's gundi; French: Goundi du Sahara; German: Langhaargundi.

physical characteristics

Head and body 6.7–9.5 in (17–24 cm); tail 1.4 in (3.5 cm); captive males averaged 6.05 oz (172 g), females 6.85 oz (194 g). Various shades of yellow, brown or buff; ears round and flat; tail bushy.

distribution

Central Sahara Desert in Algeria, northern Niger, northwestern Chad, northeastern Mali, and southwestern Libya.

habitat

Rock outcrops and mountains in desert; occurs up to 7,875 ft (2,400 m) above sea level.

behavior

Lives in rock crevices, using many temporary shelters. Diurnal; emerges to feed in early morning; peak activity one to four hours after dawn, but active for much of the day, except during hottest periods. Does not emerge in cold or wet weather; often sunbathes and grooms. Utters an infrequent chirp. In colonies, occurs at a density of 0.12/acre (0.3/ha). Lives in family groups with close social ties; females help other females during pregnancy and when giving birth.

feeding ecology and diet

Eats leaves, stems, flowers and seeds, preferring Cruciferae (especially Moricandia arvensis), Compositae, and Graminae.

reproductive biology

Young recorded March to June; females also pregnant in April–May. Estrus cycle in captive females averages 24.9 days; estrus period October to March. Litter size in captivity two to three; newborn young 2.8–3.2 in (7–8 cm), and approximately 0.7 oz (20 g). Sexually mature in eight to 12 months.

conservation status

Not globally threatened.

significance to humans

None known.

Desert gundi

Ctenodactylus vali

taxonomy

Ctenodactylus vali Thomas, 1902, north of Tripoli, Libya.

other common names

English: Sahara/Val's gundi; French: Goundi du Sahara; German: Sahara-Gundi.

physical characteristics

Head and body length is about 6.3–6.7 in (16–17 cm); tail 0.4–0.6 in (1–1.5 cm); average mass 6.2 oz (175 g). Buff to chestnut,

with whitish underparts; tail small, not bushy.

distribution

Southern Morocco, western Algeria, and northwestern Libya.

habitat

Rocky outcrops in desert, from sea level to 8,200 ft (2,500 m).

behavior

Shelters in rock crevices. Diurnal, emerging soon after dawn; activity peaks two hours after dawn, and there is often a small amount of activity for two to four hours before dusk. Sunbathes frequently. Has whistling calls. Family groups occupy territories in colonies; one colony covered approximately 26,900 ft² (2,500 m²).

feeding ecology and diet

Feeds entirely on vegetation, preferring Cruciferae (especially Eremophyton chevallieri), Compositae (especially Amberboa leucantha), and Graminae (especially Cymbopogon and Aristida); also eats acacia leaves.

reproductive biology

Semicaptive females produced young mainly in March–April, but breeding season may extend from February to June. Estrous cycle averages 23.4 days; anestrus from May to December. Gestation period 56 days; litter size one to three. Young weigh about 0.7 oz (20 g) when born; nurse for several weeks; fully grown and sexually mature in nine to 12 months. Apparently only one litter per year. Longevity five years in captivity.

conservation status

Not globally threatened. Locally threatened by hunting, human settlement and predation by domestic cats and dogs.

significance to humans

Some are hunted for food.

Resources

Books

George, Wilma. "Notes on the Ecology of Gundis." In The Biology of Histricomorph Rodents, edited by I. W. Rowlands and B. J. Weir. London: Academic Press, 1974.

George, Wilma. "Gundis." In The Encyclopedia of Mammals, edited by David Macdonald. New York: Facts on File, Inc., 2001.

Hartenberger, J. L. "The order Rodentia: Major Questions on Their Revolutionary Origin, Relationships and Superfamilial Systematics." In Evolutionary Relationships Among Rodents: A Multidisciplinary Analysis, edited by W. P. Luckett and J. L. Hartenberger. New York: Plenum Press, 1985.

Meester, J., and H. W. Setzer. The Mammals of Africa: An Identification Manual. Washington, DC, Smithsonian Institute Press, 1977.

Nowak, Ronald M. Walker's Mammals of the World. Vol. 2, 6th ed. Baltimore: Johns Hopkins University Press, 1999.

Rosevear, D. R. The Rodents of West Africa. London: The British Museum (Natural History), 1969.

Periodicals

Beintema, J. J., K. Rodewald., G. Braunitzer, J. Czelusniak, and M. Goodman. "Studies on the Phylogenetic Position of the Ctenodactylidae (Rodentia)." Molecular Biology and Evolution 8 (1991): 151–154.

George, W. "Reproduction in Female Gundis (Rodentia: Ctenodactylidae)." Journal of Zoology 185 (1978): 57–71.

Graur, D., A. Zharkikh, W. Hide, and W-H. Li. "The Biochemical Phylogeny of Guinea Pigs and Gundis, and the Paraphyly of the Order Rodentia." Comparative Biochemistry and Physiology 101 (1992): 495–498.

Wood, A. E. "The Evolution of the Rodent Family Ctenodactylidae." Journal of the Paleontological Society of India 20 (1977): 120–137.

Barry Taylor, PhD