Short-nosed elephant shrew

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Short-nosed elephant shrew
Short-nosed elephant shrew (Elephantulus brachyrhynchus)

Short-nosed elephant shrew ( Elephantulus brachyrhynchus )

Systematics
Superordinate : Afrotheria
without rank: Afroinsectiphilia
Order : Elephant jerk (Macroscelidea)
Family : Elephant jerk (Macroscelididae)
Genre : Elephant shrews ( Elephantulus )
Type : Short-nosed elephant shrew
Scientific name
Elephantulus brachyrhynchus
( A. Smith , 1836)

The short-nosed elephant shrew ( Elephantulus brachyrhynchus ), sometimes also short-nosed elephant shrew or short-nosed elephant shrew , is a type of elephant shrew . The elongated, trunk-like nose is typical, but it appears comparatively shorter than that of other members of the genus, as well as the long hind and short front legs. The species is endemic to Africa and has a large distribution area that extends from East to South Africa. Their habitat includes savannas and open forest landscapes with dense underground vegetation . There the short-nosed elephant shrew lives on the ground and is active from day to twilight. The main diet consists of insects and green parts of plants. The animals form monogamous pair relationships and use grazing areas made up of grassy areas and bushes. A litter consists of one or two young animals, whereby the number of offspring is partly influenced by the season. The population of the short-nosed elephant shrew is considered safe. The species was scientifically introduced in 1836.

description

Habitus

The short-nosed elephant shrew is one of the smaller representatives of the elephant shrews . It reaches a total length of 17.7 to 23 cm, of which 8.5 to 11.2 cm are on the tail. For animals from northern South Africa, a head-trunk length of 8.6 to 11.5 cm and a tail length of 5.7 to 9.3 cm are specified, for individuals from Uganda the corresponding values ​​for head and trunk are 10, 5 to 11.2 cm and for the tail at 8.3 to 10.2 cm. The tail has on average around 90% of the length of the rest of the body. The weight varies from 30 to 55 g. According to studies of about 30 individuals from Botswana , the average weight of males was around 44.5 g, that of females around 42.7 g. In an equally large group from northeastern South Africa , males weighed an average of 43.6 g, while females weighed 45.9 g. Typical of all elephant shrews are the large head with the elongated, trunk-like nose, which is relatively shorter compared to other species, and the short front and long rear legs. The fur is soft, and the hair on the back is up to 10 mm long. They are usually colored gray at the base, the tips have a variable shade of brown depending on the region. This results in regionally deviating back shading that is viewed as clinical . Animals from Namibia and northern Botswana are characterized by a reddish-brown back, those from South Africa by a yellowish-brown, while in turn individuals from Malawi and Mozambique appear gray-brown. In East Africa , on the other hand, animals mostly found in gray, in Zambia reddish- gray . The back fur is often interspersed with longer, black-tipped hair. The ventral side has predominantly a light gray to whitish tint. The two-colored tail is deep brown on the upper side in animals with an overall darker back coloration, in lighter-colored individuals it is more yellow-gray. The bottom is lighter than the top. At the tail end no conspicuous tufts of hair is formed, as shown when partly sympatric living Bushveld Elephant Shrew ( Elephantulus intufi occurs). On the head, whitish-gray upper lips and the equally colored, sometimes yellowish-brown appearing ring around the large eyes protrude. The ears reach a length of 19 to 24 mm and have rounded tips. Behind them there is usually a yellowish-brown stain that contrasts with the fur on the back. The undersides of the hind feet show a brown color, similar to the dryland elephant shrew and different from the black soles of the eastern cliff elephant shrew ( Elephantulus myurus ). The rear foot length varies between 27 and 34 mm.

Skull and dentition features

The skull reaches a length of 32.2 to 34.4 mm, at the zygomatic arches it is 16.8 to 18.8 mm wide. Overall, the skull has a narrow structure. The rostrum is comparatively shorter than that of other elephant shrews and is therefore more similar to the short-eared elephant shrew ( Macroscelides proboscideus ). In contrast to numerous other elephant shrews, a third molar is formed in the lower jaw , which has a cylindrical shape but only small in size. In this characteristic, the short-nosed elephant shrew resembles the dark one ( Elephantulus fuscus ) or the dark-footed elephant shrew ( Elephantulus fuscipes ). Thus, the bite has a total of 42 teeth and has the following dental formula: . Only occasionally do the rearmost (third) molars also appear in the upper jaw. In the upper row of teeth, the inner and outer incisors (I1 and I3) exceed the middle ones in height, the canine is shaped like a molar. The entire upper row of teeth is 17 mm long on average.

distribution

Distribution area (brown) of the short-nosed elephant shrew

The range of the short-nosed elephant shrew is the largest of all elephant shrews and also the elephant shrew . It occurs from northern South Africa , northern and eastern Botswana and northeastern Namibia via Swaziland , southern Mozambique , Malawi , Zambia and Zimbabwe to Angola and the south of the Democratic Republic of the Congo . In the northeast, the range extends to East Africa , where the species occurs in parts of Tanzania , Kenya and Uganda . The habitat includes the forest lands of the Zambezi as well as the adjoining areas of the Somalia-Maasai bushland and the southern and eastern rainforest-savanna mosaic landscapes. The preferred habitats are therefore savannahs with dense and tall grass vegetation, sometimes also bushes. In addition, riparian forests are also sought out by the short-nosed elephant shrew, for example in the Okavango Delta . It occurs mainly in areas with sandy to firm, but not rocky subsoil.

Studies in Zimbabwe resulted in the circulation area of the short-snouted elephant shrew a predominance of mixed woodlands with Mopane - and long threads -Pflanzengemeinschaften, moreover, also Miombo -Waldsavannen and forest and jungle areas where besides Langfäden- also Myrobalan prevail -Pflanzengesellschaften. The vegetation cover should also be dense in the dry season . The population density varies, but the species can sometimes be found locally quite frequently. In a woodland-savannah landscape on the Sengwa in northeastern Zimbabwe, only 0.2 to 0.4 individuals per hectare were identified, depending on the season. In contrast, a significantly higher population density was observed in the Mankwe nature reserve in northern South Africa. Here it was 8 to 14 individuals per hectare in open forest landscapes, but only 1 to 2 in pure grass areas. In both biotopes the population density was higher in winter than in summer. In the southern distribution area, there is some overlap with the eastern cliff and dryland elephant shrew, but the former prefers rockier habitats, the latter requires less densely vegetated areas. In the north-eastern area of ​​the occurrence there are also overlaps with the red-brown elephant shrew ( Elephantulus rufescens ).

Way of life

Territorial behavior

The short-nosed elephant shrew is mostly diurnal and twilight-active and lives on the ground, where it moves as a fast ( cursorial ) runner from four-footed to jumping. She seeks the protection of the plant cover and also withdraws under it for protection or to rest. She does not build her own burrows, but sometimes uses those of other small mammals or termite nests. The animals live in monogamous pairs and can mostly be observed together in one area. They are also very territorial. They maintain action spaces , whereby those of the tied animals often overlap. Males have larger ranges with 0.25 to 0.41 ha than females, whose territories occupy 0.17 to 0.25 ha. Investigations in the Mankwe nature reserve in northern South Africa showed that the districts include grassy areas as well as bushes and thickets, which are used depending on the various activities. The bushes into which the short-nosed elephant shrew can retreat to protect against predators or in case of bush fires are particularly important . In addition, the bushes often form the core of new action spaces, insofar as the surrounding grass areas were destroyed in a fire. In the action areas there are various activity areas, between which the short-nosed elephant shrew sometimes creates special paths and paths. However, this does not take place to the same extent as, for example, with the red-brown elephant shrew. The distances covered within a day may be rather short. Successive observations of individual individuals were made at short distances of 20 to 42 m from one another.

Fragrances from skin glands are often used for communication within the species. There are also various noises, such as a long meowing as an alarm call, which lasts up to 545 ms and reaches a frequency of 1.65 kHz. Fighting animals, on the other hand, use high-pitched squeaks. Another characteristic is the drumming of the feet, which is performed by hitting the ground with the hind feet. In the short-nosed elephant shrew, it consists of a sequence of 10 to 30 regular beats at intervals of 30 to 40 ms. Individual irregular strokes are also interspersed in this regular sequence. A new sequence of drum beats is generated at intervals of 100 to 200 ms, so that a whole series can last up to five seconds.

nutrition

The short-nosed elephant shrew is largely omnivorous. Macroscopic analyzes of stomach contents from Tanzania revealed insect remains of 47.2% , green plant material of 51.7% and seeds and fruits to be 1.1% . Most of the insect remains were ants , there were also termites and, to a very small extent, beetles . Similar results were obtained from a study of stomach contents and excrement residues in the South African Mankwe Nature Reserve, which showed about 37% insects, 60% plants and 3% seeds. Observations in Zimbabwe also show a significantly higher proportion of insects, while the proportion of plants is less than 2%. In South Africa, on the other hand, it has been shown that the short-nosed elephant shrew also consumes nectar , including from Cytinus visseri , a member of the mallow family . When looking for nectar and licking up the substance, pollen often sticks to the trunk, which is then carried on by the animals. Some of the pollen also reaches the gastrointestinal tract, where studies have shown up to 5000 pollen grains. Because of this, the short-nosed elephant shrew can also be seen as an important pollen distributor.

The body temperature is around 34.2 to 37.7 ° C. The short-nosed elephant shrew uses various options to regulate the heat balance. If there is a threat of overheating at outside temperatures of over 35 ° C, there is increased evaporation of surface water. The animals encounter low outside temperatures with extensive sunbathing, which mainly takes place in the early morning hours. The occurrence of a torpor in poor environmental conditions is not known for the species.

Reproduction

Reproduction is not seasonally bound in large areas of the distribution area, with the exception of the populations in the southern distribution area, where the offspring is likely to be born seasonally. Young animals could mostly only be observed here in the Mankwe nature reserve, for example, in the summer months (September to February). Studies in Zimbabwe showed seasonal fluctuations in the frequency of the offspring. They show that fewer young are born in the cooler parts of the year from May to August, as the food supply in the form of insects is significantly lower at this time. Here, too, the interval between two litters of a female is 90 days, while it shrinks to 60 during the warmer months of the year. A litter includes between one and two newborns. The number of newborns depends on the season and the size of the mother. Smaller females weighing less than 40 g give birth to an average of between 1.4 (cooler season) and 1.7 (warmer season) young animals. For heavier animals over 40 g, the corresponding values ​​are 1.6 to 2. The annual average number of young that are born per female is thus around 8.3, with an overall average birth rate of 1.6. It is possible that the number of young per litter is also regionally dependent, since observations indicate that an average of only 1.4 young animals were born in populations in Tanzania. The gestation period is estimated at around 50 days. In general, the newborns are well developed, completely covered in fur, and have open eyes. Their head-to-torso length is 36 to 50 mm, their weight around 8 to 9 g. Individual development is relatively quick. Life expectancy in the wild is unknown, animals lived in human care for over four years and two months.

Predators and parasites

The barn owl and the African grass owl are particularly important predators . External parasites mainly include ticks such as Rhipicephalus and Ixodes . Fleas, such as Dinopsyllus and Xenopsylla , and mites, such as Schoutedenichia , were also detected. Furthermore provides Parasubulura from the group of nematodes is an internal parasite.

Systematics

Internal systematics of elephants according to Heritage et al. 2020
 Macroscelididae  
  Macroscelidinae  
  Macroscelidini  


 Galegeeska


   

 Petrodromus


   

 Petrosaltator




   

 Macroscelides



  Elephantulini  

 Elephantulus



  Rhynchocyoninae  

 Rhynchocyon



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The short-nosed elephant shrew is a separate species from the genus of elephant shrews ( Elephantulus ), to which a total of eight other species are assigned. The elephantulus that over large parts of southern and eastern Africa are common, form part of the family of shrews (Macroscelididae) within the same order (Macroscelidea). The order comprises rather small, endemic animals occurring in Africa and is now divided into two subfamilies. The Rhynchocyoninae represent a monotypical group and contain only the proboscis dogs ( Rhynchocyon ). These represent the largest members of the elephants and predominantly inhabit densely forested habitats . The Macroscelidinae, in turn, include the elephant shrews , the proboscis ( Petrodromus ), the North African elephant shrew ( Petrosaltator ), the Somali elephant shrew ( Galegeeska ) and the species of the genus Macroscelides . These are adapted to mostly drier and more open landscapes and thus occur both in savannahs and in desert-like regions. Molecular genetic studies showed that the two subfamilies separated from each other in the Lower Oligocene about 32.8 million years ago. Within the Macroscelidinae there was greater diversification with the beginning of the Upper Oligocene around 28.5 million years ago.

Internal system of elephant shrews according to Smit et al. 2011
 Elephantulus  


 Elephantulus fuscus


   

 Elephantulus fuscipes



   



 Elephantulus rufescens


   

 Elephantulus brachyrhynchus



   

 Elephantulus rupestris


   

 Elephantulus intufi




   

 Elephantulus myurus


   

 Elephantulus edwardii


   

 Elephantulus pilicaudus






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The representation here neglects the positions of Petrodromus , Petrosaltator , Galegeeska and Macroscelides , all four of which are currently deeply embedded in Elephantulus , making the genus of elephant shrews paraphyletic .

Sir Andrew Smith
Historical representation of the short-nosed elephant shrew from 1839

From a molecular genetic point of view, the elephant shrews currently form a paraphyletic group, as Petrosaltator , Petrodromus and Macroscelides are currently still deeply embedded in the genus Elephantulus . However, the genetic studies also advocate the close relationship of a group of species common to South and East Africa. The closest relative of the short-nosed elephant shrew is therefore the red-brown elephant shrew ( Elephantulus rufescens ). The sister group of these two forms a clade consisting of the dryland elephant shrew ( Elephantulus intufi ) and the western cliff elephant shrew ( Elephantulus rupestris ). Another clade of predominantly South African species is closely related to this common family group, including the eastern cliff elephant shrew ( Elephantulus myurus ), the Cape elephant shrew ( Elephantulus edwardii ) and the Karoo cliff elephant shrew ( Elephantulus pilicaudus ), which was only newly described in 2008 . The origin of this species community may be in eastern Africa. Some of these early representatives of the elephant shrews then migrated during a phase of increasing aridization of the continent at the beginning of the Upper Miocene around 11.5 million years ago to the south and southwest and thus to the current areas of distribution. A further fragmentation of the elephant shrews resulted from a renewed desiccation of the landscapes of southern Africa in the transition from the Upper Miocene to the Pliocene around 6 million years ago. As a rule, no subspecies of the short-nosed elephant shrew are distinguished; the regional color variations that occur are considered clinical . Originally, however, the dark-footed elephant shrew ( Elephantulus fuscus ) and the dark-footed elephant shrew ( Elephantulus fuscipes ) were considered to be identical to the short-nosed elephant shrew due to the development of a third, lower molar. In addition, the trait was used to separate the genus Nasilio from Elephantulus in 1906 (and both due to the less distended tympanic membranes of Macroscelides ). However, a revision of the elephant shrew in 1968 united Nasilio with Elephantulus and at the same time recognized the dark-footed elephant shrew as an independent species, while the dark elephant shrew six years later was classified as an independent species. Subsequent investigations on allozymes and isoenzymes of elephants confirmed that the species originally assigned to Nasilio actually belong to Elephantulus . The presence of a third mandibular molar is therefore regarded as originally within the Proboscis, the reduction of which occurred several times independently in different genera. In contrast to the anatomically assumed close relationship of the short-nosed elephant shrew to the dark and dark-footed elephant shrew, the genetic investigations have so far not shown any direct relationship between the three species.

Fossil finds of the short-nosed elephant shrew are rather rare. A small amount of evidence in the form of mandibular and maxillary remains came from Kabwe and Twin Rivers. Both sites are in Zambia and date to the Middle Pistocene, but the former could also belong to the Old Pleistocene . Some uncertain finds also come from Kromdraai and Makapansgat in South Africa, but it is possible that these represent representatives of Elephantulus that are now extinct . None of the known finds has been scientifically examined in more detail so that there is still uncertainty about the actual first appearance of the short-nosed elephant shrew.

The first scientific description of the short-nosed elephant shrew was in 1836 by Andrew Smith . Smith led the species under the scientific name Macroscelides brachyrhynchus . As a type region, he indicated the region between Latakoo and the Tropic , which means Kuruman in the South African province of North Cape and Bechuanaland , today's Botswana. Like the common name , the scientific species name refers to the comparatively short form of the trunk.

Threat and protection

There are currently no known major threats to the population of the short-nosed elephant shrew. The distribution area includes the largest of all known elephants. The mostly dry grasslands only lead to a few overlaps with human settlement and economic areas. It is possible that desertification or bush cover can have an impact on the individual populations . However, it is currently not known whether there is an increasing or decreasing population trend. For these reasons, the IUCN classifies the short-nosed elephant shrew as "not endangered" ( least concern ). The species occurs in several protected areas. In human care, the short-nosed elephant shrew is rarely looked after. In the history of European zoos, there has only been one owner, London, and the animals were presented there in the first half of the 1970s.

literature

  • Stephen Heritage: Macroscelididae (Sengis). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 206-234 (p. 232) ISBN 978-84-16728-08-4
  • Mike Perrin: Elephantulus brachyrhynchus Short-snouted Sengi (Short-snouted Elephant-shrew). In: Jonathan Kingdon, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, 2013, pp. 263-265
  • Galen B. Rathbun: Elephantulus brachyrhynchus (A. Smith, 1836) - Short-snouted elephant-shrew. In: John D. Skinner and Christian T. Chimimba (Eds.): The Mammals of the Southern African Subregion. Cambridge University Press, 2005, pp. 28-29

Individual evidence

  1. ^ A b c d R. W. Yarnell and DM Scott: Notes on the ecology of the short-snouted sengi (Elephantulus brachyrhynchus) at a game ranch in North-west Province, South Africa. Afrotherian Conservation 4, 2006, pp. 2-4
  2. ↑ Erik Thorn and Julian Kerbis Peterhans (with the participation of Jonathan Baranga, Michael Huhndorf, Rainer Hutterer and Robert Kityo): Small mammals of Uganda. Bats, shrews, hedgehog, golden-moles, otter-tenrec, elephant-shrews, and hares. Bonn Zoological Monographs 55, 2009, pp. 1–164 (pp. 102–106)
  3. ↑ a b c d e f g h Galen B. Rathbun: Elephantulus brachyrhynchus (A. Smith, 1836) - Short-snouted elephant-shrew. In: John D. Skinner and Christian T. Chimimba (Eds.): The Mammals of the Southern African Subregion. Cambridge University Press, 2005, pp. 28-29
  4. ↑ a b c d e f g h i j Mike Perrin: Elephantulus brachyrhynchus Short-snouted Sengi (Short-snouted Elephant-shrew). In: Jonathan Kingdon, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, 2013, pp. 263-265
  5. ↑ a b c d G. B. Corbet and J. Hanks: A revision of the elephant-shrews, Family Macroscelididae. Bulletin of the British Museum (Natural History) Zoology 16, 1968, pp. 47-111
  6. ↑ a b c d e f g Stephen Heritage: Macroscelididae (Sengis). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 206-234 (p. 232) ISBN 978-84-16728-08-4
  7. ^ John Eric Hill: Notes on the Dentition of a Jumping Shrew (Nasilio brachyrhyncha). Journal of Mammalogy 19 (4), 1938, pp. 465-467
  8. ^ Galen B. Rathbun, David O. Ribble, Timothy O. Osborne and Seth J. Eiseb: Sengis (Elephant-shrews) in North-eastern Namibia. Journal of Namibia Scientific Society 65, 2017, pp. 69-75
  9. ↑ a b c d e B. R. Neal: The ecology and reproduction of the Short-snouted Elephant-Shrew, Elephantulus brachyrhynchus, in Zimbabwe with a review of the reproductive ecology of the genus Elephantulus. Mammal Review 25, 1995, pp. 51-60
  10. ↑ a b C. Stuart, M. Perrin, C. FitzGibbon, M. Griffin (IUCN SSC Afrotheria Specialist Group) and H. Smit (Stellenbosch University): Elephantulus brachyrhynchus. The IUCN Red List of Threatened Species. Version 2014.3. ( [1] ); last accessed on March 19, 2015
  11. ↑ AV Linzey and MH Kesner: Small mammals of a woodland-savannah ecosystem in Zimbabwe. I. Density and habitat occupancy patterns. Journal of Zoology 243, 1997, pp. 137-152
  12. ^ Richard W. Yarnell, Daniel J. Metcalfe, Nigel Dunstone, Niall Burnside, and Dawn M. Scott: The impact of fire on habitat use by the short-snouted elephant shrew (Elephantulus brachyrhynchus) in North West Province, South Africa. African Zoology 43 (1), 2008, pp. 45-52
  13. ↑ AS Faurie, ER and MR Dempster Perrin: Footdrumming patterns of southern African elephant-shrews. Mammalia 60 (4), 1996, pp. 567-576
  14. ^ A b H. Leirs, R. Verhagen, W. Verheyen and MR Perrin: The biology of Elephantulus brachyrhynchus in natural miombo woodland in Tanzania. Mammal Review 25, 1995, pp. 45-49
  15. ↑ Steven D. Johnson, Priscilla M. Burgoyne, Lawrence D. Harder and Stefan Dötterl: Evidence for pollination by small mammals in the parasitic plant Cytinus visseri (Cytinaceae). South African Journal of Botany 74 (2), 2008, p. 369
  16. ↑ Steven D. Johnson, Priscilla M. Burgoyne, Lawrence D. Harder and Stefan Dötterl: Mammal pollinators lured by the scent of a parasitic plant. Proceedings of the Royal Society B 278, 2011, pp 2303-2310
  17. ^ Mike Perrin: Comparative aspects of the metabolism and thermal biology of elephant-shrews (Macroscelidea). Mammal Review 25, 1995, pp. 61-78
  18. ↑ Colleen T. Downs and Mike R. Perrin: The thermal biology of three southern African Elephant-shrews. Journal of thermal Biology 20 (6), 1995, pp. 445-450
  19. ^ Gea Olbricht: Longevity and fecundity in sengis (Macroscelidea). Afrotherian Conservation 5, 2007, pp. 3-5
  20. ↑ LJ Fourie, JS du Toit, DJ Kok and IG Horak: Arthropod parasites of elephant-shrews, with particular reference of ticks. Mammal Review 25, 1995, pp. 31-37
  21. ↑ JP Hugot and D. Cuisance: Etude morphologique de Parasubulura gerardi Berghe & Vuylsteke, 1938 (Nematoda, Subuluroidea) parasite d'un Macroscelidae Ethiopia. Systematic Parasitology 16, 1990, pp. 67-76
  22. ↑ a b Steven Heritage, Houssein Rayaleh, Djama G. Awaleh and Galen B. Rathbun: New records of a lost species and a geographic range expansion for sengis in the Horn of Africa. PeerJ 8, 2020, p. E9652, doi: 10.7717 / peerj.9652
  23. ↑ a b c d Hanneline Adri Smit, Bettine Jansen van Vuuren, PCM O'Brien, M. Ferguson-Smith, F. Yang and TJ Robinson: Phylogenetic relationships of elephant-shrews (Afrotheria, Macroscelididae). Journal of Zoology 284, 2011, pp. 133-143
  24. ^ Mike Perrin and Galen B. Rathbun: Order Macroscelidea - Sengis (Elephant-shrews). In: Jonathan Kingdon, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, 2013, pp. 258-260
  25. ↑ Oldfield Thomas and Harold Schwann: The Rudd exploration of South Africa. V. List of mammals obtained by Mr. Grant in the North East Transvaal. Proceedings of the Zoological Society of London 1906, pp. 575-591 ( [2] )
  26. ^ J. Raman and Mike Perrin: Allozyme and isozyme variation in seven southern African Elephant-shrew species. Journal for Mammalian Science 62, 1997, pp. 108-116
  27. ^ Percy M. Butler: Macroscelidea, Insectivora and Chiroptera from the Miocene of East Africa. Palaeovertebrata 14 (3), 1984, pp. 117-200
  28. ↑ DM Avery: Early and Middle Pleistocene environments and hominid biogeography; micromammalian evidence from Kabwe, Twin Rivers, and Mumbwa Caves in central Zambia. Palaeogeography, Palaeoclimatology, Palaeoecology 189, 2003, pp. 55-69
  29. ↑ TN Pocock: Plio-Pleistocene fossil mammalian microfauna of Southern Africa - a preliminary report including description of two new fossil muroid genera (Mammalia: Rodentia). Palaeontologia Africana 26, 1987, pp. 69-91
  30. ↑ Patricia A. Holroyd: Past records of Elephantulus and Macroscelides: geographic and taxonomic issues. Afrotherian Conservation 7, 2009, pp. 3-7
  31. ↑ Patricia A. Holroyd: Macroscelidea. In: Lars Werdelin and William Joseph Sanders (eds.): Cenozoic Mammals of Africa. University of California Press, Berkeley, London, New York, 2010, pp. 89-98
  32. ^ Andrew Smith: Report of the Expedition for Exploring Central Africa. Cape Town, 1836, pp. 1–68 (p. 42) ( [3] )
  33. ^ Andrew Smith: Illustrations of the Zoology of South Africa. Mammalia. London, 1839 (Plate 13) ( [4] )
  34. ↑ Zoo animal list ( [5] ), last accessed on June 15, 2015

Web links

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