Elephant shrews

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Elephant shrews
Dryland elephant shrew (Elephantulus intufi)

Dryland elephant shrew ( Elephantulus intufi )

Systematics
Subclass : Higher mammals (Eutheria)
Superordinate : Afrotheria
without rank: Afroinsectiphilia
Order : Elephant jerk (Macroscelidea)
Family : Elephant jerk (Macroscelididae)
Genre : Elephant shrews
Scientific name
Elephantulus
Thomas & Schwann , 1906

The elephantulus ( Elephantulus ), partly trunk mice are a genus of the family of shrews (Macroscelididae) within the same order in the German (Macroscelidea). They are not related to the shrews from the order of the insectivores . The nine species of this genus live in eastern, central and southern Africa and are therefore the most widespread among the elephants. The elephant shrews are mainly adapted to dry landscapes and occur in semi-deserts , savannas and bush countries. They are all small animals, the appearance of which is characterized by a large head with a trunk-like elongated nose, thin limbs with short front and long hind legs and a long tail. They live on the ground and are known as fast runners and jumpers. Their food consists mainly of insects and only a small part of plant material. Noteworthy is the social system, which consists of monogamous couples that last a lifetime. The offspring usually consists of only one or two young animals and is raised without the assistance of the father, and maternal care is also limited. Exact information on the way of life is only available for a few species.

The first representatives of the elephant shrews were discovered and described in the early 1830s, at that time they were mostly related to the short-eared elephant shrew from the genus Macroscelides . However, differences in skull structure led to the establishment of the genus Elephantulus at the beginning of the 20th century . Today the genus is the most species-rich among the elephants, but the problem is the exact relationship of the individual representatives to each other and to other genera of the elephant. This makes it likely that the elephant shrews in their current definition do not form a uniform group. In tribal history , the elephant shrews first appeared in the Pliocene around 5 million years ago. They are often found in the fossil record , but there is a lack of meaningful studies on the individual finds. From the point of view of species protection, there are no threats to the population for the majority of species, and insufficient data are available for some.

description

Habitus

Eastern cliff elephant shrew ( Elephantulus myurus )

The elephant shrews are among the smallest members of their family, they have a total length of 20.2 to 22.8 cm in smaller species such as the short-nosed elephant shrew ( Elephantulus brachyrhynchus ) up to 23.9 to 29.2 cm in larger forms such as the western cliff elephant shrew ( Elephantulus rupestris ). The tail is usually relatively long and reaches between 67 and 130% of the length of the rest of the body. The weight is 35 to 70 g, a pronounced sexual dimorphism is not developed. Particular characteristics of the elephant shrews consist in the large head with the characteristic trunk-like elongated, very mobile nose and the lower mouth as well as in the thin limbs, in which the hind legs are significantly longer than the front legs. The fur is soft, its color varies from yellow-gray to reddish brown on the back, the underside usually appears lighter with a whitish to light gray color. Within the species there is partly a dependency of the coat color on the surrounding ground, so that the coat pattern can be viewed as clinical . In most representatives, the tail also shows a significantly darker upper side and a lighter underside. Some forms have a brush-like tuft of elongated hair at the tail end, otherwise the tail hair is differently thick. The ears become relatively large and are rounded at the tips. There is often a noticeable patch of color behind the ears. The large eyes are framed by a light eye ring, which, however, like the dark-footed elephant shrew ( Elephantulus fuscipes ) rather inconspicuous or like the western cliff elephant shrew and the Karoo cliff elephant shrew ( Elephantulus pilicaudus ) cannot be fully developed. Most species have glands on the tail to secrete scent marks, a few such as the red-brown elephant shrew ( Elephantulus rufescens ) or the dark-footed elephant shrew also on the chest area. The females have three pairs of teats . The front and rear legs each end in five rays that have claws. The four outer fingers protrude conspicuously over the innermost one ( pollex ), which only extends to the base of the next outer finger. The inner toe ( hallux ) is also shorter than the outer toe by the length of the claw. Overall, the rear foot is significantly longer than the front foot.

Skull and dentition features

The skull has a length of 32.2 to 38.7 mm and a width at the zygomatic arches of 16.8 to 21.9 mm. It is triangular in shape when viewed from above and has a bulging forehead line when viewed from the side, the rostrum appears narrow. In contrast to Macroscelides , the elephant shrews do not have a bulging tympanic bladder on the temporal bone , the volume of the middle ear is about 81 mm³ and thus only takes up about a ninth of the space compared to Macroscelides . The occiput is therefore much wider, but the entire posterior area of ​​the skull is narrower. There are three pairs of openings in the median jawbone and the palatine bone , the number and size of which correspond to those of the proboscis and the Macroscelides species, but they differ from the proboscis dogs . The lower jaw is long and slender and has high articular processes. The bit has the following dental formula : . A total of 40 to 42 teeth can therefore be formed. The additional lower third molar occurs only in the short-nosed, dark-footed and dark elephant shrews ( Elephantulus fusus ) and is rather small and round. In contrast to Macroscelides , the teeth of the front dentition are not in a closed row. The incisors are mostly small and have no sharp edges or points, and the canine is also small. The rear molars are less crowned than in Macroscelides . The length of the upper row of teeth is 16.2 to 21.9 mm.

distribution and habitat

The elephant shrews live in Africa , all known species are distributed in the east and south of the continent. Taken together, the elephant shrews are the most widespread of all elephant shrews, but the individual species are, with exceptions, regionally limited in their occurrence. All representatives are tied to rather dry landscapes. As a result, they inhabit a number of different habitats , ranging from semi-deserts to steppes and savannahs to open tree and shrubbery landscapes. Due to this specialization, they are absent in dense forests, where they are replaced by other elephants. Some of the species such as the Cape ( Elephantulus edwardii ), the Eastern cliff ( Elephantulus myurus ) and the Western cliff ( Elephantulus rupestris ) or the Karoo cliff elephant shrew, which was only described in 2008, prefers rocky to stony terrain made up of large boulders or scree fields and only has a low cover of vegetation. Others, such as the dryland ( Elephantulus intufi ), the short-nosed ( Elephantulus brachyrhynchus ), the reddish brown ( Elephantulus rufescens ), the dark ( Elephantulus fuscus ) and the dark-footed elephant shrew ( Elephantulus fuscipes ), are on the other hand fine-grained to sandy, but adapted to hard subsoil, which is mostly covered with dense vegetation. The adaptation to different substrates means that some representatives appear sympatric , but then they do not occupy the same ecological niche . In general, elephant shrews are relatively rare, but locally, depending on the seasons, they can occur in a very high population density.

Way of life

Territorial behavior

Western cliff elephant shrew ( Elephantulus rupestris )
Short-nosed elephant shrew ( Elephantulus brachyrhynchus )

The way of life has only been well studied in a few species of elephant shrews, this mainly concerns the red-brown elephant shrew ( Elephantulus rufescens ) and the eastern cliff elephant shrew ( Elephantulus myurus ), with others it is partially unexplored. All representatives of the elephant shrews are ground-dwelling and represent fast ( cursorial ) runners. They move four-footed running and jumping forward and reach speeds of up to 28 kilometers per hour. The animals are adapted to this locomotion primarily through their long and slender hind legs. It is noticeable that the metatarsus is the same length as the upper leg section, a feature that is not usually found in other small mammals such as rodents of comparable size , but is typical of numerous members of the artifacts , such as deer and some horned animals . In addition, there is the formation of a toe gait , which is also more reminiscent of artifacts than small mammals. Equipped with such good running properties and combined with their constant alertness, which results from a very well developed sense of sight , smell and hearing , elephant shrews are able to flee quickly from potential predators or other dangers.

The main activities of the animals take place during the twilight phases, but sometimes also during the day or at night. Depending on the preferred landscape, crevices or small caves as well as bushes and thickets serve as shelter; they rarely use the burrows of other animals, which then mostly come from rodents or termites . The elephant shrews use action spaces and are partly territorial, they mark their territorial boundaries with the secretions from scent glands. Within the districts, they set up a network of paths and paths that connect the individual shelters with a wide variety of feeding and resting places and are used for rapid movement. You clean these paths with swinging movements of the arms of small stones, branches, leaves and other obstacles. The social system of the elephant shrews consists of monogamous pair bonds that last a lifetime. This system has been studied particularly well in the red-brown, eastern cliff and dryland elephant shrews ( Elephantulus intufi ), and in some cases also in the short-nosed elephant shrew ( Elephantulus brachyrhynchus ). The pair bonds are not particularly strong and deep, since joint activities largely only take place during the reproductive phase. The roaming areas of the tied animals partially, and in some cases completely, overlap. Intruders are mostly driven away by the partner of the same sex. A female animal whose partner has disappeared is sometimes temporarily drawn into a polygyne connection by a neighboring, bonded male , but this only lasts until the female has found a new, unbound partner.

In addition to the exposure of scent marks, a drum drum is mainly used for intra-species communication. The animals produce the drum-like noises when their hind feet hit the ground; they consist of a sequence of regular and irregular beats of short duration that are repeated at certain time intervals. This form of communication, known as podophony , is typical for elephants. Since each type of elephant shrew has its own characteristic drum series, this has a taxonomic value. Drumming occurs mainly in stressful situations, such as territorial fights or mating rituals. It may, however, also be used interspecific, for example to inform ground-dwelling predators that they have been noticed.

Diet and thermoregulation

Dryland elephant shrew

The diet of elephant shrews consists largely of insects such as ants and termites , as well as other invertebrates . In some species, the consumption of green parts of plants, seeds and fruits has also been proven. Also for some species, such as the Cape elephant shrew ( Elephantulus edwardii ) and the short-nosed elephant shrew, nectar could also be documented in the menu; these representatives also represent important disseminators of pollen . The respective proportions of animal and vegetable food depend on the seasons, the vegetarian diet often increases when the general food supply is more diverse. The search for food takes place on the ground, under stones and bushes and is partly related to the maintenance of the path system. The nose is used for probing, the recording is made with the long, sticky tongue, which can protrude several millimeters in front of the nose.

The elephant shrews show a certain degree of heterothermia with sometimes clearly fluctuating body temperatures over the course of a day or an activity phase. For thermoregulation especially sunbathing in the morning hours or increased discharge of surface water in extreme heat used. In the rocky regions, the animals use crevices with more balanced temperatures as resting places and shelters. Species that live in desert-like climates have specially adapted kidneys to store water and concentrate urine . Due to the significantly greater temperature difference between day and night in the desert-like regions or at high altitudes, a daily torpor has formed in some species that occur there. Such rigid phases have been documented for the Cape, Eastern Cliff and Western Cliff elephant shrews ( Elephantulus rupestris ). The duration of the torpor is often linked to the outside temperature, while the body temperature drops extremely low during rigidity and reaches values ​​that are otherwise known in animals with hibernation . Rigid phases do not only occur at extreme outside temperatures, but sometimes also when there is a shortage of food.

Reproduction

Reproduction has only been well studied in a few species. As a rule, it takes place all year round, but there may be seasonal differences in the frequency of births. Only in the dryland and eastern cliff elephant shrews are reproduction seasonally restricted and take place during the warmer or wetter months. The gestation period is 50 to 57 days, one litter includes one or two young. The young weigh around 10 g and flee the nest and can walk within a very short time after birth. The boys are housed in a separate shelter; there is no paternal care. The mother animal only visits the young for a short time for daily suckling. This type of rearing of the offspring is known as the “mother absenteeism system” and is also known from other elephants. This probably means that the young remain largely odorless and are thus better protected from predators. Weaning begins as early as 25 to 30 days. Some of the offspring are driven from their common territory by their parents after they reach sexual maturity after around 50 days; they then look for their own space of action. Since females are ready to conceive again shortly after birth, the interval between two litters is around 60 to 90 days. This ensures that offspring can be born several times a year even in species with a restricted reproductive phase. Overall, however, the reproduction rate is rather low due to the long gestation period and the low number of newborns per litter. The life expectancy of an elephant shrew in the wild is often only one to two years, the highest known age of an animal in human care was more than nine years.

Systematics

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


 Galegeeska


   

 Petrodromus


   

 Petrosaltator




   

 Macroscelides



  Elephantulini  

 Elephantulus



  Rhynchocyoninae  

 Rhynchocyon



Template: Klade / Maintenance / Style

The elephantulus form a genus of the family of shrews (Macroscelididae) within the same order of shrews (Macroscelidea). The elephants are an endemic group of smaller mammals found in Africa . They include a total of six genera, which are divided into two subfamilies. The proboscis dogs ( Rhynchocyon ) are the only member of the subfamily Rhynchocyoninae , this is thus monotypical . They represent the largest representatives of the elephants, their main area of ​​distribution are predominantly forested habitats . The second subfamily, the Macroscelidinae , includes the elephant shrews as well as the trunk rat ( Petrodromus ) and the genera Macroscelides , Galegeeska and Petrosaltator . All representatives of the Macroscelidinae inhabit much drier open landscapes from savannah to desert-like regions. According to molecular genetic studies, the elimination of the elephants into the two subfamilies began in the Lower Oligocene around 32.8 million years ago, while the greater diversification of the Macroscelidinae began in the Upper Oligocene around 28.5 million years ago. Within the Macroscelidinae the elephant shrews are in the tribe of the Elephantulini, all other genera in the tribe of the Macroscelidini .

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


 Elephantulus fuscus


   

 Elephantulus fuscipes



   

 Galegeeska


   
 " Panelephantulus clade "  

 Macroscelides


   

 Petrodromus


   

 Petrosaltator




   



 Elephantulus rufescens


   

 Elephantulus brachyrhynchus



   

 Elephantulus rupestris


   

 Elephantulus intufi




   

 Elephantulus myurus


   

 Elephantulus edwardii


   

 Elephantulus pilicaudus








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The individual species of elephant shrews are sometimes difficult to distinguish externally. The sometimes strongly varying coat pattern, which is now regarded as clinical , led in the past to the description of numerous species, which are now considered synonymous with existing ones. Morphologically, the recognized species can be identified based on the presence and absence of glands in the chest area, the shape of the supratragus on the auricle , the relative size of the tympanic bladder , the number of rear molars in the lower jaw and the number, arrangement and shape of the cusps distinguish the front molars from each other. With the help of molecular genetic studies, the species could also be clearly separated from one another, but they also speak for a high variability within the genus Elephantulus and partly also within the individual species. As a result, the genetic data support the close relationship between a group of predominantly South and East African species, which had already been determined by previous studies, such as the alloenzymes and isoenzymes , but also by phylogenetic studies. The differences here lie in the details, for example in the development of a third lower molar in the short-nosed and the dark elephant shrew , which, contrary to earlier assumptions, does not suggest a closer relationship between the two species. The same applies to the hairy nasal mirror, which only occurs in the red-brown elephant shrew. It is also known from the Somali elephant shrew ( Galegeeska revoili ) and therefore cannot be seen as an expression of a closer relationship between the two representatives. However, the results of the genetic investigations did not show a conclusive position within the genus for all members of the elephant shrews, which is partly due to the limited sample material available. In addition to the dark elephant shrew, this also applies to the dark-footed elephant shrew ( Elephantulus fuscipes ).

Nine species living today are distinguished:

Originally, the North African elephant shrew ( Petrosaltator rozeti ) was also included among the elephant shrews. The distribution area of ​​this species extends from Morocco to Libya and is thus north of the Sahara , far separated from the other members of the elephant. The molecular genetic analyzes speak for a closer position of this with Petrodromus and with Macroscelides , whereby the genus Elephantulus became paraphyletic . In addition to the genetic evidence of this closer relationship between the North African elephant shrew and the proboscis, there are also morphological and anatomical indications. These include the similar structure of the penis with two laterally formed lobes near the tip or the appearance of mammary glands in male animals. Both species show similarities in general skull structure to the other elephant shrews, while the Macroscelides species differ more due to their bulging tympanic bladders. However, Macroscelides again agrees more closely with Petrodromus and the North African elephant shrew in the structure of the Basicranium , and all three representatives have an ossified nerve canal on the inner ear. Because of these similarities, the North African elephant shrew, the proboscis and the representatives of the genus Macroscelides were provisionally brought together in the common Panelephantulus clade . One possibility was a reassessment of the North African elephant shrew, a union of it with the proboscis in one genus or the merging of all three generic representatives of the Macroscelidinae in a single genus. In 2016, a group of researchers led by John P. Dumbacher decided to take the first route and relegated the North African elephant shrew to the genus Petrosaltator .

The Somali elephant shrew was also assigned to the elephant shrews. The shape is native to northeast Africa and was only known from a few museum specimens until the 21st century. It was not until 2019 that live specimens were observed in Djibouti . A genetic examination to clarify the relationship from 2011 led to an unsatisfactory result, which was mainly explained by the poorly preserved DNA samples. Only the discoveries in 2019 made further analyzes possible. They produced a closer relationship between the Somali elephant shrew, the proboscis and the North African elephant shrew. This group had split off in the Lower Miocene around 20.6 million years ago. For this reason, Steven Heritage and Houssein Rayaleh referred the Somali elephant shrew to the newly created genus Galegeeska in 2020 . They defined these by, among other things, the hairy nasal mirror , the tuft of the tail and the light eye ring marked on the back by a dark spot.

In addition to the nine recent species, two that are now extinct are recognized:

Elephantulus broomi was originally described by Robert Broom in 1937 as Elephantomys langi based on a few lower jaws and several parts of the skull from South African caves, but the following year he equated Elephantomys with Elephantulus . However, since the name langi had already been occupied in 1929 for a subspecies of the short-nosed elephant shrew, Gordon Barclay Corbet and John Hanks renamed the fossil species to Elephantulus broomi in 1968 . The species Elephantulus antiquus , also established by Broom, is based on an upper jaw from Sterkfontein .

Research history

The first representatives of the elephant shrews were discovered and described as early as the early 1830s. Until the end of the 19th century, all known species of elephant shrews were included in the genus Macroscelides . This was set up in 1829 by Andrew Smith , who assigned it to the short-eared elephant ( Macroscelides proboscideus ). In 1906, Oldfield Thomas and Harold Schwann recognized the morphological diversity of the individual species. They therefore divided up the genus Macroscelides and left only those species in this that were characterized by an inflated tympanic bladder on the temporal bone and two lower molars . This mainly affected the short-eared elephant and some representatives who are now considered synonymous with it . For Elephantulus they put the forms whose tympanic bladder was moderately large, which corresponded to the largest part of the elephant shrews known at the time. However, with Nasilio , Corbet and Hanks introduced a third genus, the members of which were distinguished from the Elephantulus representatives by the presence of three, instead of two rear, lower molars. Both authors therefore referred the short-nosed elephant shrew and the dark elephant shrew to this genus. Elephantulus and Nasilio were considered independent genera in the first half of the 20th century, and it was not until 1951 that Nasilio was designated as a subgenus of Elephantulus . In their major revision of the elephant in 1968, Gordon Barclay no longer considered Corbet and John Hanks Nasilio .

In addition to Elephantulus and Nasilio , Robert Broom introduced another genus in 1937, Elephantomys . In addition to the extinct species Elephantomys langi (now Elephantulus broomi ), he also placed the dryland elephant shrew, which he called " Elephantulus " intufi , in these . Broom saw the reference to a separate genus justified by the molar-like expression of the second premolar in the upper jaw in both species. Only a year later, Broom noticed that the western cliff elephant shrew, the type species of Elephantulus , had such a tooth and elephantomys synonymized with Elephantulus . Nevertheless, Elephantomys received the status as a subgenus of the elephant shrews in 1953, which was reversed in the revision of 1968.

Tribal history

The first representatives of the subfamily Macroscelidinae appeared in the Middle Miocene around 15 million years ago in eastern Africa. The genus Elephantulus can be detected for the first time in the Lower Pliocene and is present at numerous sites in southern and eastern Africa. The oldest known finds include those of Langebaanweg in southwestern South Africa . The finds come from river deposits and are around 5 million years old, an exact species assignment has not yet been made. The very fossil-rich cave site at Makapansgat , also in South Africa, dates back to the end of the Pliocene around 3 million years ago , from which an extensive fauna of small mammals originates, which is interpreted as the prey remains of large birds of prey . Among these are countless remains of jaws and teeth from at least 250 individuals of elephant shrews. Most of it is referred to Elephantulus antiquus , a now extinct, relatively large form that may be close to the Eastern Cliff or Cape Elephant Shrew. A smaller part, on the other hand, belongs to the now extinct species Elephantulus broomi , which is significantly smaller and possibly related to the dryland elephant shrew. Some researchers also consider both species to be identical. Both Elephantulus antiquus and Elephantulus broomi occur in the transition to the Pleistocene also at other South African sites, for example in Sterkfontein and Swartkrans , the latter has also been handed down in the Olduvai Gorge in East Africa. Since the two species were sometimes confused in the past, a reassessment is requested.

Representatives of today's species are also passed down comparatively early. The dark elephant shrew already occurs in Makapansgat in the Upper Pliocene, as well as in Swartkrans and Sterkfontein and in the Olduvai Gorge in the Lower Pleistocene . The dryland elephant shrew is again present in the South African cave sites of the Lower Pleistocene, while other representatives of the genus appear in the course of the Middle Pleistocene , such as the short-nosed elephant shrew in Kabwe and Twin Rivers in Zambia as well as the western cliff elephant shrew at Elands Bay in the southwest South Africa. The problem is that, with only a few exceptions, none of the fossil remains of the recent species were exactly described, but rather formally determined and the named species are therefore only included in the respective fauna lists. For this reason, inaccuracies or possible incorrect assignments cannot be ruled out. Other finds, on the other hand, are difficult to assign and therefore only have vague determinations, such as those from the Wonderwerk Cave in South Africa, or, as in the case of the fossil remains of the Malapa Cave , also South Africa, have to be investigated using complex methods.

Threat and protection

Most representatives of the elephant shrews are only slightly threatened in their population, which is related to their distribution in mainly dry landscapes that are little used or influenced by humans. Only in river valleys or in the vicinity of water can there be overlaps with agriculturally used areas or impairments due to settlement construction or industrialization. As a result, the majority of the species are currently “not endangered” ( least concern ). However, due to the lack of information about distribution, population size and lifestyle, the IUCN lists the dark-footed elephant shrew ( Elephantulus fuscipes ) and the dark-footed elephant shrew ( Elephantulus fuscus ) in the “ data deficient ” category. Some of these species may be at greater risk of being threatened.

literature

  • GB 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
  • 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 ISBN 978-84-16728-08-4
  • 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. 261-276
  • Ronald M. Nowak: Walker's Mammals of the World . Johns Hopkins University Press, 1999, ISBN 0-8018-5789-9
  • Galen B. Rathbun: Genus Elephantulus Thomas & Schwann, 1906. In: John D. Skinner and Christian T. Chimimba (eds.): The Mammals of the Southern African Subregion. Cambridge University Press, 2005, pp. 27-34

Individual evidence

  1. ↑ a b c d e f g h i j k l m n o 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. 261-276
  2. ↑ a b c d e f g h 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
  3. ^ Galen B. Rathbun: Genus Elephantulus Thomas & Schwann, 1906. In: John D. Skinner and Christian T. Chimimba (eds.): The Mammals of the Southern African Subregion. Cambridge University Press, 2005, pp. 27-34
  4. ↑ a b c d e f g h i j 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 ISBN 978-84-16728-08-4
  5. ^ Matthew J. Mason: Structure and function of the mammalian middle ear. I: Large middle ears in small desert mammals. Journal of Anatomy 2015, doi: 10.1111 / joa.12313
  6. ↑ Jan Ihlau, Friederike Kachel and Ulrich Zeller: Graphical description of the ventral side of a sengi's (Macroscelides proboscideus) skull. Afrotherian Conservation 4, 2006, pp. 11-12
  7. ↑ a b Fabiana Panchetti, Massimiliano Scalici, Giuseppe Maria Carpaneto and Giancarlo Gibertini: Shape and size variations in the cranium of elephant-shrews: a morphometric contribution to a phylogenetic debate. Zoomorphology 127, 2008, pp. 69-82
  8. ↑ a b c d 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
  9. ↑ a b c d e f Galen B. Rathbun: Why is there discordant diversity in sengi (Mammalia: Afrotheria: Macroscelidea) taxonomy and ecology? African Journal of Ecology 47, 2009, pp. 1-13
  10. ↑ Barry G. Lovegrove and Metobor O. Mowoe: The evolution of micro-cursoriality in mammals. The Journal of Experimental Biology 217, 2014, pp. 1316-1325
  11. ^ Galen B. Rathbun: The social structure and ecology of Elephant-shrews. Zeitschrift für Tierpsychologie Beiheft 20 (Advances in Behavioral Research), 1979, pp. 1-76
  12. ^ David O. Ribble and Michael R. Perrin: Social organization of the Eastern Rock Elephant-shrew (Elephantulus myurus): the evidence for mate guarding. Belgian Journal of Zoology 135 (suppl.), 2005, pp. 167-173
  13. ^ Galen B. Rathbun and CD Rathbun: Social structure of the bushveld sengi (Elephantulus intufi) in Namibia and the evolution of monogamy in the Macroscelidea. Journal of Zoology 269, 2006, pp. 391-399
  14. ↑ a b 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
  15. ^ 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
  16. ↑ AS Faurie, ER and MR Dempster Perrin: Footdrumming patterns of southern African elephant-shrews. Mammalia 60 (4), 1996, pp. 567-576
  17. ↑ Petra Wester: Sticky snack for sengis: The Cape rock elephant-shrew, Elephantulus edwardii (Macroscelidea), as a pollinator of the Pagoda lily, Whiteheadia bifolia (Hyacinthaceae). Natural Sciences 97, 2010, pp. 1107–1112
  18. ↑ 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
  19. ^ Mike Perrin: Comparative aspects of the metabolism and thermal biology of elephant-shrews (Macroscelidea). Mammal Review 25, 1995, pp. 61-78
  20. ↑ CT Downs: Renal structure, and the effect of an insectivorous diet on urine composition of Southern African Elephant-Shrew species (Macroscelidea). Mammalia 60 (4), 1996, pp. 577-589
  21. ^ BG Lovegrove, J. Raman and MR Perrin: Heterothermy in elephant shrews, Elephantulus spp. (Macroscelidea): daily torpor or hibernation. Journal of Comparative Physiology 171, 2001, pp. 1-10
  22. ↑ Rebecca Oelkrug, Carola W. Meyer, Gerhard Heldmaier and Nomakwezi Mzilikazi: Seasonal changes in thermogenesis of a free-ranging afrotherian small mammal, the Western rock elephant shrew (Elephantulus rupestris). Journal of Comparative Physiology B 182, 2012, pp. 715-727
  23. ↑ Fritz Geiser and Nomakwezi Mzilikazi: Does torpor of elephant shrew differentiate from other hetherothermic did of mammals? Journal of Mammalogy 92 (2), 2011, pp. 452-459
  24. ↑ EG Sauer: On the social behavior of the short-eared elephant shrew Macroscelides proboscideus. Zeitschrift für Mammaliankunde 38, 1973, pp. 65-97
  25. ^ Gea Olbricht: Longevity and fecundity in sengis (Macroscelidea). Afrotherian Conservation 5, 2007, pp. 3-5
  26. ^ CJ van der Horst: Some remarks on the biology of reproduction in the female of Elephantulus, the holy animal of Set. Transactions of the Royal Society of South Africa 31 (2), 1946, pp. 181-199
  27. ↑ a b c d e 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
  28. ↑ a b c d e f 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
  29. ^ A b G. B. Corbet: A cladistic look at classification within the subfamily Macroscelidinae based upon morphology. Mammal Review 25 (1/2), 1995, pp. 15-17
  30. ↑ a b 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
  31. ↑ Hanneline Adri Smit, Terence J. Robinson, Johan Watson and Bettine Jansen van Vuuren: A New Species of Elephant-shrew (Afrotheria: Macroscelidea: Elephantulus) from South Africa. Journal of Mammalogy 89 (5), 2008, pp. 1257-1268
  32. ^ Matjaž Kutner, Laura J. May-Collado and Ingi Agnarsson: Phylogeny and conservation priorities of afrotherian mammals (Afrotheria, Mammalia). Zoologica Scripta 40, 2011, pp. 1-15
  33. ^ PF Woodall: The penis of elephant shrews (Mammalia: Macroscelididae). Journal of Zoology 237, 1995, pp. 399-410
  34. ^ Gea Olbricht and William T. Stanley: The topographic distribution of the penis and mammary glands in sengis (Macroscelidea) and its usefulness for taxonomic distinctions. Zoosystematics and Evolution 85 (2), 2009, 297-303
  35. ^ A b Massimiliano Scalici and Fabiana Panchetti: Morphological cranial diversity contributes to phylogeny in soft-furred sengis (Afrotheria, Macroscelidea). Zoology 114, 2011, pp. 85-94
  36. ↑ a b c Julien Benoit, Nick Crumpton, Samuel Merigeaud and Rodolphe Tabuce: Petrosal and Bony Labyrinth Morphology Supports Paraphyly of Elephantulus Within Macroscelididae (Mammalia, Afrotheria). Journal of Mammalian Evolution 21, 2014, pp. 173-193
  37. ↑ John P. Dumbacher, Elizabeth J. Carlen and Galen B. Rathbun: Petrosaltator gen. Nov., A new genus replacement for the North African sengi Elephantulus rozeti (Macroscelidea; Macroscelididae). Zootaxa 4136 (3), 2016, pp. 567-579
  38. ^ A b Robert Broom: On some new Pleistocene mammals from limestone caves of the Transvaal. South African Journal of Science 33, 1937, pp. 750-768
  39. ^ Robert Broom: Some South African Pliocene and Pleistocene Mammals. Annals of the Transvaal Museum 21, 1948, pp. 1-38
  40. ↑ 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 ( [1] )
  41. ^ Robert Broom: Note on the premolars of the elephant shrews. Annals of the Transvaal Museum 19 (2), 1938, pp. 251-252
  42. ^ QB Hendey: Palaeoecology of the late Tertiary fossil occurrences in 'E' Quarry, Langebaanweg, South Africa, and a reinterpretation of their geological context. Annals of the South African Museum 84 (1), 1981, pp. 1–104 (p. 50)
  43. ^ D. Margaret Avery: An assessment of the Lower Pleistocene micromammalian fauna from Swartkrans Members 1-3, Gauteng, South Africa. Geobios 31 (3), 1998, pp. 393-414
  44. ↑ 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
  45. ↑ Percy M. Butler: Insectivora and Chiroptera. In: Vincent J. Maglio and HBS Cooke (Eds.): 11I Evolution of African Mammals. Harvard University Press, 1978, pp. 56-68
  46. ↑ 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
  47. ^ Thalassa Matthews, Christiane Denys and John E. Parkington: The palaeoecology of the micromammals from the late middle Pleistocene site of Hoedjiespunt 1 (Cape Province, South Africa). Journal of Human Evolution 49, 2005, pp. 432-451
  48. ↑ Patricia A. Holroyd: Past records of Elephantulus and Macroscelides: geographic and taxonomic issues. Afrotherian Conservation 7, 2009, pp. 3-7
  49. ↑ DM Avery: Pleistocene micromammals from Wonderwerk Cave, South Africa: practical issues. Journal of Archaeological Science 34, 2007, pp. 613-625
  50. ↑ Aurore Val, Kristian J. Carlson, Christine Steininger, Job M. Kibii, Cecil Churms, Brian F. Kuhn and Lee R. Berger: 3 D techniques and fossil identification: An elephant shrew hemi-mandible from the Malapa site. South African Journal of Sciences 107 (11/12), 2011, pp. 1–5 ( [2] )
  51. ↑ Degree of endangerment of the individual species in the IUCN Red List of Threatened Species .

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