Fat-tailed gerbil

Fat-tailed gerbil
Fat-tailed gerbil ( Pachyuromys duprasi )
Systematics
Superfamily : Mice-like (Muroidea) Family : Long-tailed mice (Muridae) Subfamily : Gerbils (Gerbillinae) Tribe : Desmodilliscini Genre : Pachyuromys Type : Fat-tailed gerbil
Scientific name of the  genus
Pachyuromys
Lataste , 1880
Scientific name of the  species
Pachyuromys duprasi
Lataste , 1880

The fat-tailed gerbil ( Pachyuromys duprasi ) is a type of gerbil and the only type of the genus Pachyuromys . Mostly it is assigned to the "real gerbils" ( Gerbillini ), but recent studies indicate a basal position within the gerbil and a closer relationship with the brewer gerbil . Named after Hippolyte Dupras, it was described by Fernand Lataste in 1880 . It is not endangered and is kept as a test and pet animal.

Like the brewer gerbil, it is one of the most specialized desert-dwelling gerbils. Its distribution area is the northern edge of the Sahara and it inhabits sparsely vegetated semi-deserts on different soils, including sand and gravel. In addition to leaves and seeds, it feeds on insects and happily accepts living crickets in human care.

With its body about nine and a half to twelve centimeters long without a tail and a tail that is only half a body length, it looks like a larger version of the brewer's gerbil. It differs from most other gerbils by its almost rounded and plump body, the rather short legs, the short, thickened tail and the short and pointed snout, which is more reminiscent of a hamster than a gerbil.

In contrast to all other gerbils, it is able to store fat in its tail for times when there is little food. This can swell like a club, especially when cared for by humans. Her timpani bubbles , which are used in the open landscape to hear particularly far-reaching, deep tones, are particularly large and therefore sensitive. They protrude far beyond the back of the skull and considerably exceed the brain volume.

Body features

Exterior

The fat-tailed gerbil has a stocky body. The auricles are rounded, as long as the fur and strongly arched inward. The legs are short and strong, the tarsus short. There are four toes on the front paws and five on the hind paws. The claws are curved and pointed, those of the front paws are clearly elongated. The soles of the front and rear paws are partially hairy. The pads including the big toe pad are similar to those of the real gerbils and racing rats .

The fur is long and fluffy, on top of a light yellowish gray to brownish yellowish brown. The underside, the front legs and the rear paws are white. The area around the mouth is partially pigmented. The auricles are also pigmented and sparsely hairy. There is a tuft of long, cinnamon-colored hair at their front approach, and there is a white spot behind each auricle.

The tail is almost hairless, two-colored, on the upper side like the sides of the body, white on the underside and has no tassel . It serves as a storage organ for fats , which can be metabolized from adipose tissue , and thus for water . A thin, wrinkled tail indicates malnutrition or illness .

According to Kingdon (1997) the head-torso length of the fat-tailed gerbil is 105 to 135 millimeters, the tail length 45 to 61 millimeters and the body weight 30 to 65 grams. Niethammer (1988) gives the head-torso length as 95 to 120 millimeters, the tail length as 50 to 65 millimeters and the body weight as 20 to 45 grams. The body dimensions of male and female adult animals are almost the same.

skull

The skull of the fat-tailed gerbil is elongated and triangular when viewed from above due to the enormously inflated tympanic bladders. The bony ridge ( crista supraorbitalis ) above the eye socket is poorly developed and does not reach forward to the rear edge of the tearbone , the lateral ridge ( crista parietalis ) on the parietal bone is inconspicuous. The parietal bone is narrow and triangular with the tip pointing backwards. The anterior surface of the tympanic bladder lies in front of the foramen ovale of the sphenoid bone and its posterior margin protrudes well beyond the posterior margin of the supraoccipital and paroccipital of the occiput . The fossa subarcuata of the temporal bone is small, the fossa mastoidea other hand, very large and rearwardly through an association of the descending bone extension of the supraoccipital and the ascending portion of the processus suprameaticus of the temporal bone sealed. The external auditory canal is swollen and touches the zygomatic process of the temporal bone. There is a secondary eardrum ( tympanum accessorium ). The supraoccipital and basioccipital of the occiput are narrow and constricted by the inflated tympanic sacs. The paroccipital process is elongated, thin and fused with the wall of the tympanic bladder. The parapterygoid fossa of the sphenoid bone is deep, constricted by the tympanic bladder, and has a large opening in the upper wall. The rounded front edge of the zygomatic plate does not extend to the seam between the intermaxillary bone and the upper jaw . In adult animals, the bone sutures are fused.

The tympanic membranes are completely inflated and are the largest of all gerbils. They penetrate through the arch of the lateral semicircular canal ( canalis semicircularis lateralis ) into the wart part and fill it completely. As a result, the wart part, which is unique in its internal structure, only has one atrium, the antrum tympanomastoideum . The vertical, massive bone wall in front of the mastoid cavity and separating it from the tympanic cavity probably corresponds to the septum ( septum mastoideum ) of the mastoid cavity in the "higher gerbils" ( Rhombomyina ), the brewer gerbils and the "lower gerbils" ( Gerbillina ). A septum bordering the mastoid cavity below resembles and corresponds, because it includes the lateral semicircular canal, probably to the septum ( septum tympanomastoideum ) between the tympanic cavity and mastoid cavity in the "real gerbils" ( Gerbillini ). It would therefore remain connected to the outer instead of the upper wall of the tympanic bladder along the seam between the tympanic bone and the wart part, so that the wart part neither forms a lower atrium ( antrum mastoideum ventralis ) nor a corresponding septum ( septum mastoideum ventralis ). The posterior cell ( cellula mastoidea posterior ) of the mastoid process opens like all "real gerbils" to the mastoid cavity and the corresponding septum connects the posterior semicircular canal ( canalis semicircularis posterior ) with the wall of the wart part.

The transformation of the lower jaw resembles to a certain extent that of the most highly developed grass-eating gerbils and voles , the gerbil and the field mice . The horizontal part is shortened and the angular process is pointed, the ascending branches, on the other hand, are broadened with a steep front edge. Since the fat-tailed gerbil is a generalized seed eater and also feeds on invertebrates to a considerable extent , this cannot be an adjustment to its diet. Another explanation could be the overly enlarged tympanic bladders that push the back of the lower jaw forward. However, this does not explain why the ascending branches are very wide, whereas in other gerbils with enlarged tympanic bladders they became narrower.

denture

1	·	0	·	0	·	3	=  16
1	·	0	·	0	·	3

Tooth formula of the fat-tailed gerbil

The teeth of the fat-tailed gerbil, which are typical for gerbils, have one incisor tooth and three molars in each half of the jaw. Canines and molars are absent, and the total number of teeth is 16. The incisors are white, sharply curved, and the upper incisors are furrowed in front.

The molars are rooted and of the further developed bunodont , semilophodont type with almost symmetrically arranged main cusps and without additional side cusps. The first upper and lower molar teeth show cusps in very young animals, which develop into ridges in older young animals. No cusps are indicated in the second upper and lower molar. The third molar is simple and has no lateral indentations . The melting pattern is more similar to that of the Sundevall racing rat than that of the actual gerbil . In adult animals, the tooth strips are worn out.

Habitat and way of life

habitat

The habitat of the fat-tailed gerbil is limited to the hammada , i.e. deserts with solid ground . It inhabits sandy areas with vegetation and occasionally rocky areas, and in Egypt it seems to prefer sparsely vegetated areas over the more fertile coastal deserts of the Mediterranean. In Libya, it is also most common in the transitional deserts that run roughly parallel to the lush coastal plains. Their population density is assumed to be fluctuating.

In a secluded, sandy depression east of Bir Shafarzin, a specimen was caught in populations of the goosefoot Anabasis articulata . Another specimen comes from a barren gravel area 26 kilometers northwest of Cairo. The location of the type specimen, Bir Victoria, is a small, shallow, sandy depression that is sparsely overgrown with the solitary wormwood Artemisia monosperma and in places with the henbane Hyoscyamus muticus .

In Egypt it shares its habitat with the Sundevall racing rat , the small Egyptian gerbil , the Anderson gerbil , the pale gerbil , probably with the great Egyptian gerbil and the small gerbil .

Way of life

The fat-tailed gerbil's way of life is nocturnal and it becomes active at dusk. Their construction in Egypt is simple and dug up to a meter deep in hard sand. In Algeria, on the other hand, she lives in very complex structures, which, however, may not have been dug herself. Since it roams on a large scale, it could occupy the burrows of other species. The majority of authors describe them as solitary . Others mention that she lives in small groups, presumably a female with several hatchlings. If the food supply is particularly good, it may lead to a loose merger; if the food supply is poor, it falls into a brief hibernation in winter .

Their diet in nature is not fully known. The vegetable diet includes seeds and stems from, among others, Anabasis articulata and Artemisia monosperma . In addition, it feeds on insects and possibly snails . Setzer (1957) suggested the consumption of land snails in Libya and in the Libyan desert of Egypt Osborn and Helmy (1980) found gnawed snail shells in their habitat . Fruits are brought to the building from a distance of up to two kilometers and stored there. In laboratory animal husbandry, she accepts grain , minced meat , cheese , milk , lettuce and alfalfa and has a preference for live crickets . The predators include eagle owls and barn owls , in whose crests Niethammer (1988) found the skull of the fat-tailed gerbil. Adaptations against predators could be their burrowing way of life as well as the light fur color that camouflages on the desert floor.

In the ecosystem , the fat-tailed gerbil may be important as an aerator for the desert floor due to its burrowing way of life. It likely serves as important prey for birds and is believed to have a significant impact on the populations of the insects and plants on which it feeds.

Reproduction and Life Expectancy

Nothing is known about the reproduction of the fat-tailed gerbil in nature. Flower (1932) reported litters with three to five, seven and nine pups born in human care in April, May, July, October and November. According to Hayssen and coworkers (1993), however, she throws in human care all year round, the sexual cycle lasts about seven days on average, the gestation period is 19 to 24 days, the litter size is between one and nine young animals, usually three to five, weaning occurs after 29 days and one male reached sexual maturity at two months of age. According to Baker and Yahnke (2004) she throws up to three times a year and males and females become sexually mature at three to six months. The young are nestled and naked, blind and helpless when they are born.

Jones (1982) reported an age of four years and five months for a specimen kept in human care. According to Baker and Yahnke (2004), it can reach an age of five to seven years. Above all, the wear and tear of the molars has an age-limiting effect.

Behavior and senses

The fat-tailed gerbil is rather sluggish and overslept most of the day. She often sleeps stretched out on her back. If she is touched while she is sleeping, it takes up to half a minute for her to react. Osborn and Helmy (1980) describe them as the most meek of the Egyptian rodents. It never bites or makes little effort to escape when picked up. According to Sistermann (2006), however, there are also very aggressive specimens that react to every approach with biting. Flower (1932) observed cannibalism in human care and females can eat their young.

Arguing animals prefer to bite their tails and squeak loudly. The injuries caused can lead to infection and loss of the tail. In harmless quarrels, the animals drum their front paws on and squeak. For mating behavior includes a ritualized combat, in which the animals stand on its hind legs with his front paws wrestle while squealing. If the female is not in a mating mood, she hurls earth at the male with her hind legs. During the rearing of the young, the female often drives the male out of the common nest

The males of the fat-tailed gerbil have scent glands on their belly, which they use to mark their territory . In addition to the sense of smell , the sense of touch and sight are important, and the sense of hearing is also important.

Distribution and existence

The fat-tailed gerbil is only known from the Holocene of North Africa. Its distribution area is the north of the Sahara from Morocco via Algeria , Tunisia and Libya to Egypt . It also occurs occasionally in Western Sahara and Mauritania . In Egypt, it is common in the Libyan Desert along the coastal desert of the Mediterranean, especially on its southern edge. Their distribution area extends there from the area south of as-Sallum to the west side of the Nile Delta , including the Wadi an-Natrun and the west side of the Nile valley south to Abu Roasch .

The International Union for Conservation of Nature IUCN classified the fat-tailed gerbil in 1996, 2004 and 2008 as not endangered (“least concern”). This is justified with the wide distribution, the presumably high population and the improbability of a significant decline in the population. It is common, the stocks are stable, and it is not exposed to any significant threats. Measures to protect the species of the fat-tailed gerbil were not taken. It is not known whether it occurs in protected areas.

Systematics and nomenclature

External system

The systematic position of pachyuromys within the gerbils is uncertain . It is the only species of the genus Pachyuromys and is mostly assigned to the "real gerbils" ( Gerbillini ) (Pawlinow and coworkers, 1990; McKenna and Bell, 1997; Pawlinow, 2003; Musser and Carleton, 2005).

Molecular genetic studies of mitochondrial cytochrome - b - and 12S - rRNA genes by Chevret and Dobigny (2005), however, they combine with the Brewers gerbil and place both basal to the other gerbils. In investigations using the maximum likelihood method , the closed community of descent with a bootstrap value of more than 90 percent received strong support and was always determined to be basal to the other gerbils. However, the genetic distance between the two species is 20.1 percent and is thus in the range of the values ​​determined for the other tribes, so that Chevret and Dobigny consider a classification within a new taxon in the rank of a tribe to be justified. Measurements using the relaxed molecular clock revealed a split of the two species 9.69 million years ago with a standard deviation of 1.68 million years and a split from the other gerbils 11.72 million years ago with a standard deviation of 1.82 million years.

However, these results contradict morphological investigations. Both species are very peculiar and have no known inferred similarities . According to Pawlinow (2008), the type of inflation of the wart part speaks for a placement within the "real gerbils" ( Gerbillini ), albeit with a certain restriction. The further development of the wart part is clearly a derived peculiarity of the species. It indicates that it belongs to the basal fanning of this tribe and justifies its status as a sub-tribus Pachyuromyina.

According to Tong (1989), the fat-tailed gerbil belongs to the "higher gerbils" ( Rhombomyina ) and is the sister species of the Przewalski racing rat . This hypothesis is based on similarities in the construction of the tympanic membranes and the molar crowns. So Tong assumes a change in position of the septum ( septum tympanomastoideum ) between the tympanic cavity and mastoid cavity in the "higher gerbils" towards the septum delimiting the mastoid cavity below. According to Pavlinov, however, the similarities are only superficial and the change in position of the partition is unlikely according to the principle of economy . Instead, the different positions of the partition speak for a sister group relationship with the "higher gerbils". In addition, the molars of the fat-tailed gerbil are clearly of the more developed bunodont type, while those of the "higher gerbils" are to be assigned to the prismatic type. In particular, the molars of the Przewalski racing rat are among the most specialized among the “higher gerbils” and have nothing in common with those of the fat-tailed gerbil. However, karyological examinations support membership of the "higher gerbils".

In one of the first phylogenetic schemes developed for gerbils , Petter (1959) arranged the fat-tailed gerbil together with the Somali gerbil and a few other gerbils without further developed molars as a basal arrangement within the gerbil . However, due to the diversity of the morphology of gerbils known today, Pawlinow does not consider this approach to be justified.

Huet (1881), Thomas (1882) and Sclater (1899) assigned the short-eared gerbil as Pachyuromys auricularis to the genus Pachyuromys . Lay (1972) also considers them to be closely related to the fat-tailed gerbil because of the similarities in the wart part, which has only one cavity. According to Pavlinov, however, this is a parallel development , because the tympanic bladder caused the warts part to expand along different penetration routes.

Internal system

Three subspecies of the fat-tailed gerbil can be distinguished:

• Pachyuromys duprasi faroulti Thomas, 1920 from western Algeria,
• Pachyuromys duprasi duprasi Lataste, 1880 from northern Algeria as well
• Pachyuromys duprasi natronensis de Winton, 1903 from Egypt.

The three forms show only minor deviations in essential features of both the skull and the exterior and, according to Thomas (1920), should only be viewed as local subspecies. The forms duprasi and natronensis appear to be most closely related.

This results in the following cladogram of the subspecies:

Fat-tailed gerbil	NN  Pachyuromys duprasi duprasi     Pachyuromys duprasi natronensis     Pachyuromys duprasi faroulti

nomenclature

The type specimen of Laghouat in the Sahara nordalgerischen described Fernand Lataste on 15 November 1880 as Pachyuromys duprasi , the type species of the new genus Pachyuromys . The generic name Pachyuromys is derived from ancient Greek παχυς pachys ' thick, fat ', ουρα oura ' tail ' and μυς mys ' mouse ', the specific epithet duprasi by Hippolyte Dupras . The Subertribus Pachyuromyina was described by Igor Jakowlewitsch Pawlinow in 1982.

Subspecies

Pachyuromys duprasi faroulti

The fur color of the upper side of Pachyuromys duprasi faroulti is significantly darker than that of the other two forms and is gray-brown to brownish-yellowish brown. On the sides of the body, it is sharply delimited from the underside by a wide, light cinnamon-colored band. This is not very pronounced on the cheeks and flanks, but noticeable on the trunk. The ears are brownish-yellowish white and the outer edge is brown. The fine hairs of the tail are pale cinnamon in color. The front and rear paws, on the other hand, are white as in the nominate form. The body size and essential characteristics including the skull and the dentition also resemble the nominate form.

The head-trunk length of the type specimen is 105 millimeters, the tail length 62 millimeters, the rear foot length 23 millimeters, the greatest skull length along the central axis 33.2 millimeters, the greatest skull length diagonally to the back of the tympanic membrane 36.5 millimeters, the condyloincisive length 30, 7 millimeters, the diagonal length of the tympanic membranes 17 millimeters, the bimeatal width 20.5 millimeters and the length of the upper molar row 5 millimeters.

During Walter Rothschild's expedition to western Algeria in 1913, a young fat-tailed gerbil whose coat color was darker than expected was collected near Aïn Séfra in the western Algerian plateau at an altitude of 1100 meters. Victor Faroult collected three adult animals from the same region in the summer of 1918 near Mécheria, 100 kilometers northwest of Aïn Séfra, also at an altitude of 1,100 meters. The bellows were given to Rothschild, who left them to Oldfield Thomas for examination. Following Rothschild's suggestion for naming, Thomas described the type specimen on June 15, 1920 as Pachyuromys duprasi faroulti . It bears inventory numbers 20, 3, 1, 1 of the British Museum .

Pachyuromys duprasi duprasi

The more or less ocher-brownish-yellow fur color on the upper side is comparatively pale in the nominate form Pachyuromys duprasi duprasi and gradually merges into the white of the underside on the sides of the body. The edge of the ears is not darker. The coat color corresponds to that of other gerbils in the same area a light desert shade.

Pachyuromys duprasi natronensis

Locations of Pachyuromys duprasi natronensis in Egypt to Osborn and Helmy (1980)

Pachyuromys duprasi natronensis is paler than the other two forms. The coat color is more similar to the color of the Sundevall racing rat than the Libyan racing rat . The very pale, yellowish brown-yellow coat color on the upper side gradually merges into the white of the underside on the sides of the body. The edge of the ears is not darker. The hair on the top is light cinnamon with black tips and a dark gray base. On the side of the body there is a narrow, light cinnamon-colored stripe, which extends along the hind leg almost to the heel but not along the front leg. The area around the eye is colored like the side of the body, and there is sometimes a dull spot over the eye. The spot behind the auricle is small and a white spot on the trunk is missing. There may be individual deviations in the shade and hair with black tips may be present or absent on the sides of the body.

A specimen from Bir Victoria ( 30 ° 24 ′  N , 30 ° 37 ′  E ) on the way from the Egyptian Nile to Wadi an-Natrun was described by William Edward de Winton on August 25, 1903 as Pachyuromys dupresi [sic] natronensis . Osborn and Helmy (1980) studied 14 most caught alive or excavated specimens: in addition to the type specimen from Bir Victoria three copies of Khatatba ( 30 ° 23 '  N , 30 ° 50'  O ), two additional copies of Bir Victoria, a specimen of a site 1.6 kilometers east of it, three specimens from Wadi an-Natrun ( 30 ° 25 ′  N , 30 ° 13 ′  E ), one specimen from kilometer 163 of the desert road from Cairo to Alexandria , one specimen from kilometer 26 of the same road a specimen from Abd El-Mawla ( 30 ° 33 '  N , 29 ° 13'  O ) and a copy of a location 25.6 kilometers east of Bir Shafarzin ( 31 ° 19 '  N , 24 ° 53'  O ). Setter (1952, 1963) and Hoogstraal (1963) to call other findings in Kom Hamada ( 30 ° 46 '  N , 30 ° 42'  O ), at a location between Kom Hamada and Bir Victoria, wherein Khatatba in Bir Victoria, at kilometer 179 of the desert road from Cairo to Alexandria, at kilometer 17 the same road in Abar El-Gong ( 31 ° 19 '  N , 26 ° 53'  O ) 36 kilometers west of Marsa Matruh in Qatta ( 30 ° 13 '  N , 30 ° 58 ′  E ), at a site west of Abu Ghalib ( 30 ° 16 ′  N , 30 ° 54 ′  E ), at a site west of Abu Roasch ( 30 ° 2 ′  N , 31 ° 6 ′  E ), at one site near Cairo ( 30 ° 3 ′  N , 31 ° 15 ′  E ) and at kilometer 10.5 on the desert road from Cairo to Alexandria.

Body and skull measurements of four adult Pachyuromys duprasi natronensis

Dimensions in millimeters	Average	Range of values
Head to torso length	108.3	93-121
Tail length	58.2	55-62
Ratio of tail length to head-trunk length in percent	54.4	47.9-66.6
Hind foot length	23.3	22-24
Ear length	14.0	12-16
Body weight in grams	36.5	22.0-44.6
Occipitonasal length	34.9	32.4-36.5
zygomatic width	19.3	17.5-20.2
Interorbital width	6.2	5.8-6.4
Nasal length	13.0	11.7-13.8
Length of the incisor holes	6.4	6.2-6.8
Length of the upper molar row measured on the tooth sockets	5.2	4.8-5.7
Rostral width	4.9	4.4-5.0
Height of the skull including the tympanic sacs	14.7	14.2-15.0

1. ↑ The information relates to three copies.

Fat-tailed gerbil and human

German trivial names are "Dickschwanzmaus" (Piechocki, 1969; Sauermost and Freudig, 2003), "Fettschwanzmaus" (Niethammer, 1988) and "Fettschwanz-Germaus" and "Dickschwanz-Germaus" (Sistermann, 2006).

The fat-tailed gerbil is kept as an experimental animal. After the house guinea pig, she was one of the first rodents to be found to have a vaginal plug . Héron-Royer (1881) suggested that this is formed in the vagina before copulation and pulled out or loosened by hooks on the penis . Blanchard examined tissue samples of the vaginal plugs collected by Héron-Royer and found a central part with a large number of flagellated sperm and a peripheral part formed from hardened cervical mucus . According to Lataste (1882), however, the penis is smooth and without hooks. As with other rodents, the vaginal plug forms immediately after copulation and ensures that the sperm do not flow out of the vagina. Lataste mentioned a case in which a structure similar to the vaginal plug was found without prior copulation.

In addition, the fat-tailed gerbil is kept as a pet . Couples often have no offspring for no apparent reason.

Web links

Commons : Fat-tailed gerbil ( Pachyuromys duprasi )  - Collection of images, videos and audio files

• Pachyuromys duprasi onthe IUCN Red List of Threatened Species . (with distribution map)
• Fat-tailed gerbil at Das Mäuseasyl (with information on keeping)

literature

Further and used literature:

• Ralf Sistermann: The fat-tailed gerbil ( Pachyuromys duprasi ). Biology, keeping and breeding of an unusual gerbil . In: Rodentia. Small mammal specialist magazine . No. 32 , 2006, ISSN  1617-6170 , p. 35-37 .

Used literature:

• Glover Morrill Allen: A checklist of African Mammals . In: Bulletin of the Museum of Comparative Zoology, Harvard College . tape 83 , 1939, ISSN  0027-4100 , pp. 1–763 ( full text ).
• Stéphane Aulagnier, Laurent Granjon: Pachyuromys duprasi . In: IUCN 2008 (Ed.): 2008 IUCN Red List of Threatened Species . 2008 ( full text ).
• Sheunna Barker, Chris Yahnke: Pachyuromys duprasi . In: Animal Diversity Web . ( Full text ).
• Pascale Chevret, Gauthier Dobigny: Systematics and evolution of the subfamily Gerbillinae (Mammalia, Rodentia, Muridae) . In: Molecular Phylogenetics and Evolution . tape 35 , no. 3 , 2005, ISSN  1055-7903 , p. 674-688 , doi : 10.1016 / j.ympev.2005.01.001 .
• Richard Hoath: A Field Guide to the Mammals of Egypt . The American University in Cairo Press, Cairo / New York 2003, ISBN 977-424-809-0 .
• Frederik Anna Jentink: Mammalia . In: Zoological Station Naples (ed.): Annual zoological report for 1880. IV. Department: Vertebrata . Wilhelm Engelmann, Leipzig 1881, p. 244-260 ( full text ).
• Jonathan Kingdon: The Kingdon Field Guide to African Mammals . A&C Black Publishers, London 2007, ISBN 978-0-7136-6513-0 (first edition: 1997, reprint of the original edition from 1997).
• Malcolm C. McKenna, Susan K. Bell: Classification of Mammals Above the Species Level . Columbia University Press, New York 1997, ISBN 0-231-11012-X .
• Guy G. Musser, Michael D. Carleton: Superfamily Muroidea . In: Don E. Wilson, DeeAnn M. Reeder (Eds.): Mammal Species of the World . A Taxonomic and Geographic Reference . 3. Edition. Johns Hopkins University Press, Baltimore 2005, ISBN 0-8018-8221-4 , pp. 894-1531 ( full text ).
• Jochen Niethammer: Wühler . In: Grzimek's Encyclopedia Mammals. tape 5 , p. 206–265 (eleven-volume licensed edition of the original edition from 1988).
• Ronald M. Nowak: Walker's Mammals of the World . 6th edition. Johns Hopkins University Press, Baltimore / London 1999, ISBN 0-8018-5789-9 .
• Dale J. Osborn, Ibrahim Helmy: The contemporary land mammals of Egypt (including Sinai) . In: Fieldiana: Zoology, New Series . No. 5 , 1980, ISSN  0015-0754 , pp. 1-579 ( full text ).
• Igor Jakowlewitsch Pawlinow: [Systematics of the mammals of the world] . Moscow State University, Moscow 2003 ( full text - Russian: Систематика современных млекопитающих .).
• Igor Jakowlewitsch Pawlinow: A review of phylogeny and classification of Gerbillinae (Mammalia: Rodentia) . In: Soologicheskie issledovanija . No. 9 , 2008, ISSN  1025-532X , p. 1-68 .
• Rudolf Piechocki: Family Wühler . In: Irenäus Eibl-Eibesfeldt, Martin Eisentraut, Hans-Albrecht Freye, Bernhard Grzimek, Heini Hediger, Dietrich Heinemann, Helmut Hemmer, Adriaan Kortlandt, Hans Krieg, Erna Mohr, Rudolf Piechocki, Urs Rahm, Everard J. Slijper, Erich Thenius ( Ed.): Grzimeks animal life . Encyclopedia of the Animal Kingdom. Eleventh volume. Mammals 2 . Kindler-Verlag, Zurich 1969, p. 301-344 .
• August Rauber: Ontogeny . In: Zoological station in Naples (ed.): Annual zoological report for 1882. IV. Department: Vertebrata . Wilhelm Engelmann, Leipzig 1884, p. 84-142 ( full text ).
• Rolf Sauermost, Doris Freudig and others (ed.): Lexicon of biology. In fourteen volumes . tape 11 : Phallaceae to resistance breeding . Spectrum Akademischer Verlag, Heidelberg 2003, ISBN 3-8274-0336-7 .
• William Lutley Sclater: Descriptive list of the rodents of South Africa . In: Annals of the South African Museum . tape 1 , no. 2 , 1899, ISSN  0303-2515 , p. 181-239 ( full text ).
• Charles R. Stockard, George Nicholas Papanicolaou: The vaginal closure membrane, copulation, and the vaginal plug in the guinea-pig, with further considerations of the oestrous rhythm . In: Biological Bulletin, Marine Biological Laboratory . tape 37 , no. 4 , 1919 ( full text ).
• Oldfield Thomas: A new fat-tailed gerbil ( Pachyuromys ) from western Algeria . In: Novitates Zoologicae. A Journal of Zoology in connection with the Tring Museum . tape 27 , 1920, p. 313-314 ( full text ).

Literature used indirectly:

• Martin Daly: Of Libyan jirds and fat sand rats . In: Natural History . tape 88 , 1979, pp. 64-71 .
• William Edward de Winton: Novitates Zoologicae. A Journal of Zoology in connection with the Tring Museum . tape 10 , 1903, ISSN  0950-7655 .
• David L. Harrison: The Mammals of Arabia. tape 3 : Lagomorpha, Rodentia . Ernest Benn, London 1972, ISBN 0-510-39952-5 .
• Virginia Hayssen, Ari Van Tienhoven, Ans Van Tienhoven: Asdell's Patterns of Mammalian Reproduction. A Compendium of Species Specific Data . Comstock / Cornell University Press, Ithaca 1993, ISBN 0-8014-1753-8 .
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Individual evidence

1. ↑ Pavlinow, 2008 (p. 9).
2. ↑ ^{a b c d e f} Niethammer, 1988 (p. 257, p. 264)
3. ↑ ^{a b c} Sistermann, 2006 (p. 35).
4. ↑ ^{a b} Piechocki, 1969 (p. 341)
5. ↑ ^{a b c} Nowak, 1999 (p. 1453).
6. ↑ ^{a b} Lataste, 1880 (pp. 313-315). → Quoted in: Jentink, 1881 ( p. 258 ).
7. ↑ ^{a b c d e f} Kingdon, 1997 (p. 196).
8. ↑ ^{a b c d e} Osborn and Helmy, 1980 ( p. 221 ).
9. ↑ ^{a b} Sistermann, 2006 (pp. 35–36).
10. ↑ ^{a b c} Osborn and Helmy, 1980 ( p. 223 ).
11. ↑ Osborn and Helmy, 1980 ( pp. 221-223 , Fig. 68 ).
12. ↑ Pawlinow, 2008 (p. 48, p. 19, Fig. 7g).
13. ↑ Pawlinow, 2008 (pp. 48–49, Fig. 2e).
14. ↑ ^{a b c} Osborn and Helmy, 1980 ( p. 223 , fig. 59 ).
15. ↑ Pavlinow, 2008 (p. 14, p. 48).
16. ↑ Daly, 1979. → Quoted in: Nowak, 1999 (p. 1453).
17. ↑ ^{a b c d e f g} Hoath, 2003 (pp. 186-187).
18. ^ Ranck, 1968 (p. 157). → Quoted in: Osborn and Helmy, 1980 ( p. 226 ).
19. ↑ Osborn and Helmy, 1980 ( pp. 224-226 ).
20. ↑ ^{a b c d e f g} Osborn and Helmy, 1980 ( p. 226 ).
21. ↑ ^{a b} Petter, 1961. → Quoted in: Osborn and Helmy, 1980 ( p. 226 ).
22. ↑ Sistermann, 2006 (p. 36).
23. ^ Setzer, 1957 (p. 60). → Quoted in: Osborn and Helmy, 1980 ( p. 226 ).
24. ↑ ^{a b c d e} Barker and Yahnke, 2004 ( full text ).
25. ↑ ^{a b} Flower, 1932. → Quoted in: Osborn and Helmy, 1980 ( p. 226 ).
26. ↑ Hayssen, 1993. → Quoted in: Nowak, 1999 (p. 1453).
27. ^ Jones, 1982. → Quoted in: Nowak, 1999 (p. 1453).
28. ^ Harrison, 1972. → Quoted in: Sistermann, 2006 (p. 36).
29. ↑ ^{a b} Sistermann, 2006 (p. 37).
30. ↑ ^{a b} McKenna and Bell, 1997 (p. 160).
31. ↑ ^{a b c d e} Musser and Carleton, 2005 ( " Pachyuromys duprasi " ( Memento of the original from March 15, 2009 in the Internet Archive ) Info: The archive link has been inserted automatically and has not yet been checked. Please check the original and archive link according to instructions and then remove this note. p. 1240). 
32. ↑ ^{a b} Aulagnier and Granjon, 2008 ( data set and distribution map ).
33. ↑ Pawlinow and colleagues, 1990. → Quoted in: Pawlinow, 2008 (pp. 31–32).
34. ↑ Pavlinow, 2003 ( full text ).
35. ^ Chevret and Dobigny, 2005 (p. 683).
36. ↑ Chevret and Dobigny, 2005 (Table 5).
37. ↑ ^{a b c d} Pawlinow, 2008 (p. 49).
38. ^ Pavlinow, 2008 (p. 43).
39. ↑ Pavlinow, 2008 (p. 49, Fig. 10b).
40. ↑ Tong, 1989. → Quoted in: Pawlinow, 2008 (p. 49, Fig. 10c).
41. ^ Pavlinow, 2008 (p. 29).
42. ↑ Petter, 1959. → Quoted in: Pawlinow, 2008 (p. 49, Fig. 10a).
43. ↑ Huet, 1881 (p. 339). → Quoted in: Sclater, 1899 ( p. 194 ).
44. ↑ Thomas, 1882 (p. 265). → Quoted in: Sclater, 1899 ( p. 194 ).
45. ^ Sclater, 1899 ( p. 194 ).
46. ↑ Lay, 1972. → Quoted in: Pawlinow, 2008 (p. 49).
47. ↑ ^{a b c d e f g} Thomas, 1920 ( pp. 313-314 ).
48. ↑ Lataste, 1880 (p. 313). → Quoted in: Allen, 1939 ( p. 330 ).
49. ↑ biodiversitylibrary.org
50. ↑ Pavlinow, 1982 (p. 30). → Quoted in: McKenna and Bell, 1997 (p. 160).
51. ↑ Osborn and Helmy, 1980 ( Fig. 66 ).
52. ↑ ^{a b} Osborn and Helmy, 1980 ( p. 224 ).
53. ↑ Setzer, 1963. → Quoted in: Osborn and Helmy, 1980 ( p. 223 ).
54. ↑ de Winton, 1903 (p. 285, plate 8, fig. 3). → Quoted in: Allen, 1939 ( p. 330 ).
55. ^ Setzer, 1952; Setzer, 1963; Hoogstraal, 1963. → Quoted in: Osborn and Helmy, 1980 ( p. 224 ).
56. ^ Osborn and Helmy, 1980 ( Tab. 24 ).
57. ↑ Sauermost and joyful, 2003 ( "gerbils" p 487).
58. ↑ Stockard and Papanicolaou, 1919 ( p. 230 ).
59. ↑ Héron-Royer, 1881 ( pp. 453-459 , pp. 469-472 ). → Quoted in: Stockard and Papanicolaou, 1919 ( p. 230 ).
60. ↑ Lataste, 1882 ( pp. 235-239 , pp. 258-262 ). → Quoted in: Rauber, 1884 ( p. 94 )
61. ↑ Lataste, 1882 ( pp. 235-239 , pp. 258-262 ). → Quoted in: Stockard and Papanicolaou, 1919 ( p. 230 ).