Tenreks

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Tenreks
Lesser hedgehog trek (Echinops telfairi)

Lesser hedgehog trek ( Echinops telfairi )

Systematics
Class : Mammals (mammalia)
Subclass : Higher mammals (Eutheria)
Superordinate : Afrotheria
without rank: Afroinsectiphilia
Order : Tenrecus (Afrosoricida)
Family : Tenreks
Scientific name
Tenrecidae
Gray , 1821

The Tenreks or Tanreks (Tenrecidae) are a family of mammals living in Madagascar . The group is heterogeneous gestaltlich and among others includes hedgehog - shrew - and otter-like representative. The mammals, which are not too large, are mainly nocturnal, live largely solitary and feed on invertebrates . They have adapted to different ways of life. Among the tenreks there are digging, ground-dwelling, tree-climbing and aquatic forms. As a result, the animals use various ecological niches that had remained vacant due to a lack of competition among mammals in Madagascar. At the same time, this diversity reduces intra-species competition and has led to more than thirty species, some of which differ significantly in their physique due to their different way of life. Overall, the tenreks are a prime example of adaptive radiation . In addition, the individual tenre maps sometimes have properties that are also rather unusual for other higher mammals . These include the strongly fluctuating body temperature and the occurrence of a torpor under tropical conditions, the ability of some hedgehog tenreks to stridulate with the spines of their fur , as well as the sometimes extremely short generation succession and the high number of newborns , especially in the great tenrek .

The first mention of tenreks by Western naturalists and travelers was in the middle of the 17th century. Around 100 years later, several forms were known, but they were all spiky-haired tenreks. Initially, the tenreks were therefore related to the European hedgehogs. The establishment as an independent group prevailed in the transition from the 18th to the 19th century. The characteristic appearance and the partly insectivorous diet were the reason that the tenreks were classified in the large, but not uniform group of insectivores (lipotyphla) in the course of the 19th and 20th centuries . At the end of the 20th century, however, molecular genetic studies showed that they form a separate group of mammals together with the gold mole rats (Chrysochloridae): the tenre-kitties (Afrosoricida). This group is in turn related to other originally African mammals, which are summarized under the name Afrotheria .

The origin of the Tenreks is so far unknown. There are no fossils in Madagascar that date back to the Holocene . There are also only a few remains of continental Africa, the oldest being from the Eocene of Southwest Africa. Further finds came to light in eastern Africa and date to the Lower Miocene . The group probably developed on the continent, and the ancestors of the Malagasy Tenreks arrived on the island about forty to twenty million years ago. However, it has so far been controversial among experts whether Madagascar was reached in one or in several settlement waves. Today's tenreks are adapted to both the humid rainforests of the eastern and the drier landscapes of the western part of the island. Almost a fifth of today's species are more or less threatened.

features

Habitus

Real rice burrower ( Oryzoryctes hova ), a representative of the shrew- like tenreks with soft fur
Black-headedrek ( Hemicentetes nigriceps ), a representative of the hedgehog-like tenreks with bristly fur

The tenreks are a diverse group of smaller mammals. The head-trunk length varies considerably, it reaches 4.7 to 6.4 cm in the smallest species such as the dwarf lesser rek ( Microgale parvula ) or the gnome lesser rek ( Microgale pusilla ), in the largest species, the great tenrek , it is 26, 5 to 39.0 cm. The weight fluctuates accordingly from 2.6 to 2400 g. Most species do not have any particular sexual dimorphism . Outwardly, two types of shape can be distinguished. The smaller species such as the oryzorictinae (Oryzorictinae) and Erdtenreks (Geogalinae) have a spitzmaus- , sometimes otter-like appearance with soft fur and an elongated head. The tail length of these is variable, there are species whose tail is only a third of the body length, in turn it is more than twice the length of the rest of the body in others. The second group of forms, the Igeltenreks (Tenrecinae) is characterized by a hedgehog-like appearance. Their tail forms only a short stub, their fur consists of bristles and spines. In general, the color of the tenreks' fur varies from gray to brown to black, with the underside usually being lighter in color. A conspicuous pattern of the fur is only developed in the striped duck treks ( Hemicentetes ). The hands and feet of the Tenreks each end in five toes, with the exception of the four-toed rice burrower ( Oryzorictes tetradactylus ), which has only four-pronged hands. The legs show no noticeable difference in length, the thumb and big toe are not opposable . The head looks comparatively large and, like the Great Tenrek, can make up a third of the body's length. The muzzle is generally long and flexible. The ear length varies depending on the ecological adaptation of the animals between comparatively large to small, the eyes usually remain small. Noticeably long vibrissae appear on the face .

Skull and dentition features

Great Tenrec skull ( Tenrec ecaudatus )

The skull of the Tenreks is generally elongated and narrow and flat. It has an elongated, often cylindrically shaped and more or less pointed rostrum at the front . The brain skull, on the other hand, is wider, with the greatest width usually occurring at the joint pits for the lower jaw. Especially in the hedgehog trek, the rostrum is significantly longer than the posterior section of the skull and can take up to 60% and more of the length of the skull. In the travel treks, however, the rostrum is shorter and makes up around half the length of the skull. The most striking feature is the non-closed zygomatic arch , which occurs in numerous insectivorous mammals. The area behind the eyes is often clearly indented and elongated. In the general morphology, the skulls of the Tenreks vary only slightly, in detail they show a high variety, which reflects their partly different adaptations. The rostrum can be very narrow like the striped tenrek or wider like the large tenrek or the large hedgehog tenrek. A crest is also formed differently and occurs, for example, in the great tenrek, but is absent in the striped tenrek. The brain skull is also curved or flattened to varying degrees and may be equipped with additional bones as muscle attachment points. In the embryonic stage, the skulls of all tenreks largely resemble those of the small tenreks ( microgale ).

The bit is made up of 32 to 40 teeth along the dental formula is: . A common feature of all tenreks is the absence of the first premolar . With regard to the front teeth, the individual species and genera differ significantly from each other. In principle, the incisors are equipped with a pointed crown, the appearance of individual additional cusps in front of and behind the main tip varies between the individual species. The canine tooth can resemble the incisor teeth, i.e. incisiviform and very small, such as in the small or striped duck or, as in the rice burrower and the large tenrek, more like a typical caninus ( caniniform ), in the latter it protrudes over the incisors. The arrangement of the front teeth up to and including the premolars also shows great differences, as many species have different diastemas (interdental spaces). In the structure of the premolars and the molars , the tenreks differ less clearly from each other. They are also characterized by pointed cusps, of which the molars each have three ( tritubercular or tricuspid ). The three main cusps include, based on the maxillary teeth, the paraconus, the metaconus and the protoconus. Paraconus and metaconus are close together, the protoconus is reduced in size, not as strong as in the golden mole , but stronger than in the otter shrew . There is also a V-shaped shear bar (ectoloph) on the chewing surface. Because of these characteristics, the tenrek molars can be regarded as typical zalambdodonts . Sometimes there is a cingulum on the tongue side, a bulge of tooth enamel.

Skeletal features

Skeleton of the great hedgehog trek ( Setifer setosus )

As with all Afrotheria , the spine of the Tenreks is characterized by an increased number of vertebrae (thoracic and lumbar vertebrae). It consists of 7 cervical, 15 to 19 chest, 4 to 7 lumbar, 2 to 3 sacrum and 8 to 52 caudal vertebrae. The hedgehog tails have the shortest tails with only 8 to 10 vertebrae, which results in a very short tail stub. The small long-tailed Kleintenrek ( Microgale longicaudata ), on the other hand, has the highest number of tail vertebrae among today's mammals, in addition to the long-tailed pangolin; its tail is often more than twice the length of the rest of the body.

In general, the Tenreks show a matching skeleton structure, common to all is the presence of a collarbone and the tibia and fibula that are fused together in the lower area . In addition, as in many other higher mammals , the upper ankle is only formed by the tibia and ankle bone. However, the upper ankle joint is configured somewhat differently in relation to the position of the ankles in the hedgehog treks than in the travel treks. This leads to the fact that the former keep their feet rotated outwards in the resting position, while the latter are oriented parallel to the body axis. In addition, the sometimes quite different ways of life of the Tenreks are reflected in different variations in the structure of the musculoskeletal system. These are mainly visible on the front limbs, while the rear ones are less clearly affected. The shoulder blade of burrowing representatives such as the rice burrowers ( Oryzorictes ) or the striped duck treks ( Hemicentetes ) tends to be long and narrow, whereas in climbing forms such as the small hedgehog ( Echinops ) it is short and wide. In addition, ground-burrowing tenreks have a short and wide humerus with a rather laterally pressed joint head, the shape of which restricts rotating movements. The lower joint end (elbow joint) spreads strongly laterally, which corresponds to an also clearly extended olecranon , the upper articular process of the ulna . This is where the forearm muscles come into play. In tree-dwelling species, on the other hand, the humerus is rather long and narrow, the joint head with its round shape is designed for wide rotating movements and the joints of the elbow are not so noticeably stretched. The hand and foot skeleton can be characterized by short and wide bones as in burrowing species or by longer and narrower ones as in climbing species. The former also have individual adhesions of the carpal bones , which reduces and stabilizes the mobility of the hand. In addition, the central beam is often elongated.

Soft tissue anatomy

The tenreks have five types of muscles in the front muzzle, which attach to the zygomatic bone and the anterior eye region and, with the help of tendons, extend to the tip of the nose. They allow the animals to move their elongated nose in a variety of ways. Parallel to the adaptations of the skeleton to the different ways of life, there were also changes in the muscles. While ground-dwelling animals retained a relatively original arrangement of the muscles, the semimembranosus muscle of the hind leg developed enormously in the water-dwelling water-dwelling and the burrowing striped- duck , which supports the knee when bending the leg and thus ensures powerful movement in the water or when Dig. Something similar happened on the arms, where some muscles like the teres major and the triceps brachii muscles increased in burrowing animals like the striped duck and the rice burrowers . At the same time, there are differences between the two genera in the arm and shoulder muscles, for example in the size ratio of the triceps brachii muscle to the supraspinatus muscle . The adaptations to the different ways of life within the subfamilies of the Tenreks were probably made independently of one another, which led to slightly varying modes of movement.

A noticeable feature of the tenreks is the cloaca , a common outlet for the sexual, digestive and excretory organs. The cloaca has the shape of a bowl, the entrance to the digestive tract is regulated by a sphincter muscle . There is no appendix in the digestive tract . In many species, the testes are located in the abdominal cavity ; in the Kleintenreks and the rice burrowers, they are shifted forward into the pelvis . The size of the paired testicles varies between 0.07 and 0.56 g, which corresponds to 0.8 to 2% of the body mass. The penis is very thin and extremely long, with the small long-tailed small tenrek it reaches a good 40% and with the large tenrek a good 70% of the head-trunk length, penis spines are not developed. The uterus of the females is always two-horned ( uterus bicornis ), the utero-vagina canal appears striking, which resembles a long, thin and spiral-shaped tube and is thus adapted to the shape of the penis. In contrast to the otter shrews, the tenreks have a haemochorial placenta .

The brain is one of the most simply built of all higher mammals . The weight is 420 to 2490 mg, which on average corresponds to the brain size of the shrews, but is significantly less than that of the approximately equal-sized elephants . The neocortex and striatum are very small, but the olfactory bulb is enlarged.

distribution and habitat

The tenreks are endemic to Madagascar today , only the great tenrek occurs on some smaller neighboring islands. The animals inhabit a wide variety of habitats in Madagascar, including the moist rainforests of the eastern and central parts of the island, the dry forests of the western and the thorn bush savannas of the southwest. Most, especially smaller, representatives occur in more closely defined habitats. There are three different ecotypes:

The tenreks can be found both at sea level and in high mountain areas up to 2500 m. More than one species occurs in many habitats . Up to 18 species have been detected on the Tsaratanana massif in the north, while there are still 16 species in the forest area of ​​Tsinjoarivo. The frequent occurrence together speaks for a strong ecological and trophic differentiation within the family.

Way of life

Territorial and social behavior

Large tenrek in a hollowed-out tree as a shelter
Little hedgehog climbing in the branches
Actual striped duck trek ( Hemicentetes semispinosus ) with erect spines as a defensive act

The way of life has only been well researched in very few species, this is especially true of the representatives of the hedgehog trek . With the exception of the Erdtenreks and members of the genus Nesogale, only sparse information is available for the smaller forms . In general, the Tenreks show a varied spectrum of adaptations to the most varied of lifestyles, which have arisen due to the original absence of ecological competitors such as shrews , hedgehogs or moles . A distinction can be made between different basic types: These include the underground burrowing forms such as the striped ducklings , the rice burrowers , the Erdtenreks or a few short-tailed small-tails. As a rule, the animals with short ears, small eyes and long, wide claws are adapted to the soil-burrowing way of life. There are also tree-climbing forms such as the small hedgehog trek , the representatives of the genus Nesogale or numerous long-tailed small tenreks. All climbing forms have short, strongly curved claws as an adaptation. Intermediate in between are the generally ground-dwelling forms, such as the large tenrek , the large hedgehog or small tenrek with a medium-length tail. The only water-dwelling form is the water tenrek , which has webbed feet. However, there is often an overlap between the individual basic types. For example, the water trek also creates underground structures, and numerous bottom-dwelling small treks and the large hedgehog trek can also climb.

The Tenreks are generally solitary, temporary pairings mostly only occur during the reproductive phase. An exception is the real striped duck , which lives in colonies from several generations during the mating season. The activity times of the Tenreks are mostly limited to the night, during the day they retreat to self-dug earthworks, tree hollows or other shelters. A bimodal distribution of activity, which mostly skips the midnight hours, is striking in the hedgehog treks , which have been better investigated . The animals move on the ground mainly in the cloister . Climbing tenreks are rather slow, the strongly curved claws are used like crampons, with the long-tailed small tenreks the long tail partially functions as a grasping organ . Orientation is largely based on the sense of smell . In unknown or confusing terrain, the tenreks can also emit simple echolocation calls. These are generated by clicking the tongue in a frequency range from 5 to 17  kHz . This is known from almost all hedgehog tenreks and from Dobsonian small tenreks . A special form of communication occurs with some hedgehog treks. These can use their spines to make rattling noises, which is known as stridulation . In the case of the streaked tenreks, a special stridulation organ is formed on the rear back, which consists of specially shaped spines. This also occurs in the large tenrek, but only in young animals. The little hedgehog also stridulates occasionally, with the spines responsible for this being regularly distributed over the back.

The tenreks' territorial behavior has hardly been researched. According to studies in the Ankarafantsika National Park , the Great Igeltenrek uses comparatively large action areas there , which can take up up to 13.7 hectares. When conspecifics meet, the animals sniff each other on different body parts. Aggressive behavior leads to erecting of the spines in the hedgehog tenreks, especially those in the neck area, and to typical thrusting head movements, sometimes combined with biting attacks. When threatened, the mouth is often opened, the small and the large hedgehog trek can curl up in a ball in case of danger. Little is known about secretion markings , as a rule the entrances to the burrows are marked with feces . In particularly stressful situations, Tenreks have a whitish secretion from their eyes. This is obviously a body reaction that resembles the human "eye tears".

Diet and energy balance

In keeping with their diverse lifestyles, the Tenreks' diet is also relatively varied. The main diet consists primarily of invertebrates , especially insects and earthworms . The water trek also eats crustaceans . The striped tenreks, on the other hand, specialize almost exclusively in earthworms. Some species have also been shown to kill small vertebrates or eat carrion. In addition, plant material such as fruits are consumed to a lesser extent. For many of the smaller representatives from the group of the Reistenreks , the exact diet has not been adequately researched. Here, however, isotope analyzes indicate a predominance of animal food, the composition of which varies depending on the preferred way of life. When searching for food, the nose is mainly used. The animals ingest their food with their mouths, there is hardly any manipulation with their front feet.

Most tenreks cannot regulate their body temperature to the same extent as most of the other higher mammals . Exceptions here are the small tenreks, whose body temperature is obviously more stable, but too few species have been investigated so far. In the rest of the Tenreks, the resting body temperature largely adapts to the outside temperature and is therefore subject to fluctuations. During the activity phase at night, more stable conditions are often established over several hours; in addition, the body temperature in females fluctuates less strongly in the reproductive phase. In many cases, the more stable states are accompanied by an increased metabolic rate , which is otherwise much lower than would be expected in mammals of comparable size. The Tenreks are therefore sometimes viewed as "protoendothermic" or "basoendothermic". The onset of a gate (rigid state) in unfavorable external conditions is known for several types . Such periods of rigidity can occur briefly during the southern summer from November to May, but become more extensive during the southern winter from June to October. Sometimes the animals spend this dry and relatively low-nutrient period in the torpor, a period of up to seven months is documented for the great hedgehog trek, and even up to nine months for the great tenrek. In addition to such a generally energy-saving way of life under tropical conditions, the advantage of such long periods of rest is probably that the animals can hardly be captured by predators outside of the reproductive period . This in turn results in a higher life expectancy. The tenreks' plastic behavior in terms of their energy balance and metabolism can be seen as the result of the development on a rather isolated island with little fluctuating environmental conditions and a manageable number of homoiothermal competitors. Closely related groups in continental Africa, who live under more extreme conditions, do not have as widely varying body functions. The independent increase in body temperature after a daily rest period or torpor is possibly supported by brown adipose tissue . At least in the small hedgehog trek, this could be demonstrated in smaller amounts on the neck and in larger amounts on the reproductive organs.

Reproduction

Reproduction has only been studied comparatively well in the hedgehog treks, the Erdtenreks and the representatives of the genus Nesogale . In some species, such as the Great Tenrek, polyovulation occurs, in which several egg cells are deposited in the follicle . It is noteworthy that the follicles do not form a Graafian follicle, as studies on the Great Tenrek, the Small Igeltenrek and the Striped Tenrek suggest. Rather, the granulosa cells swell , so that both the theca and the follicular epithelium become juicy. The egg cell is fertilized in the follicle, for which purpose the sperm penetrate the theca. Only then does ovulation occur . The fertilized egg slowly penetrates towards the surface of the follicle. The process is triggered by the swelling pressure of the granules. With regard to the character of the follicle, the fertilization that takes place in it and the ovulation that takes place afterwards, the tenreks differ significantly from the other higher mammals.

Family group of the Great Tenrec

As far as is known, the reproductive phase is limited to the southern summer and is therefore seasonal. It therefore falls in the wet and nutritious season. The male's advertising ritual consists of sniffing various parts of the female's body. The gestation period is usually between 50 and 70 days, it varies little between species, but can vary considerably within individual species. Accordingly, individual torpor phases influence the duration of the gestation period, as in the case of the earth tenrek or the great tenrek, as well as the respective weather. In the case of the small hedgehog trek, on the other hand, the gestation times become shorter the later the pregnancy occurs in the reproductive period. Overall, the question of influencing factors has not yet been clarified. The number of pairs of teats in the females varies between three and four in the small tenrek up to 14 in the large tenrek . Accordingly, the litter sizes are between one and up to 32 in the case of the Great Tenrek, the latter being the largest value of all mammals. In general, the soft-haired smaller tenreks have smaller litter sizes than the prickly hedgehog tenreks.

Newborn tenreks are born as nestlings and are blind as well as naked, the individual development is comparatively fast. There are striking differences in the individual phases of individual development between the prickly hedgehog treks and the soft-haired smaller species. The eyes of the hedgehog treks open after nine to 14 days, while this process lasts up to four weeks for the smaller forms. Usually the young are fully grown after one to two months. The overall short development phase and the ability of some species to become pregnant again during rearing means that females can have several litters during one reproduction phase. According to previous studies, neither the gestation period nor the individual development of the Tenreks is generally dependent on the body weight of an individual / species, which differs from the circumstances in other higher mammals. This suggests that some elements in family biology are controlled phylogenetically rather than constitutionally.

Systematics

External system

Internal systematics of Afrotheria according to Kuntner et al. 2011
 Afrotheria  
  Paenungulata  

 Sirenia (manatees)


   

 Hyracoidea (hyrax)


   

 Proboscidea ( proboscidea )




  Afroinsectiphilia  

 Tubulidentata (aardvark)


  Afroinsectivora  

 Macroscelidea (elephant)


  Afrosoricida  

 Chrysochloridae (golden mole)


  Tenrecomorpha  

 Potamogalidae (Otter shrews)


   

 Tenrecidae (Tenreks)







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The Tenreks are a family from the order of the Tenrekartigen (Afrosoricida), which also contains the otter shrews (Potamogalidae) and the golden mole (Chrysochloridae). The closest relatives of the Tenreks are the otter shrews, the close relationship to one another is expressed by the superordinate taxon of the Tenrecomorpha within the Afrosoricida. The Tenre-kartigen in turn belong to the superordinate order of Afrotheria , one of the four main lines within the higher mammals . The Afrotheria represent a kinship of animal groups predominantly native to Africa or from there. Their togetherness is based primarily on molecular genetic studies less on anatomical similarities. Within the Afrotheria there are two larger lines, on the one hand the Paenungulata and on the other hand the Afroinsectiphilia . The former include today's elephants , hyrax and manatees , the group has long been considered a common unit of descent. The latter line combines not only the tenre karts but also the elephant and the aardvark . The more precise relationships between the last three groups are under discussion. Some genetic studies advocate a sister group relationship between Afrosoricida and elephants, both groups are then combined as Afroinsectivora. Others, in turn, see the tenre cardigans in a sister group position to all other Afrotheria, while the elephants form a closer relationship with the aardvark. According to the molecular genetic investigations, the Afrotheria formed in the Upper Cretaceous 90.4 to 80.9 million years ago, their split into the two main groups followed about 15 million years later. The Afrosoricida then appeared shortly before the Cretaceous-Tertiary boundary about 68 million years ago. Around ten million later, the gold mole separated from the common line with the tenreks and the otter shrews.

The extinct Plesiorycteropus is also closely related to the Tenreks . Originally, a closer relationship to the aardvark was considered, which is also expressed in the common German name "Madagascan aardvark". However, later anatomical analyzes showed no close proximity to the aardvark; rather, numerous characteristics between the two that were regarded as coincident turned out to be an adaptation to a burrowing way of life. Based on this, Ross DE MacPhee outsourced Plesiorycteropus to his own order of Bibymalagasia. Genetic analyzes from 2013 have now shown that Plesiorycteropus is close to the Tenreks, the animals should therefore be regarded as giant tenrecs ("giant tenreks ").

Internal system

Internal systematics of the Tenreks according to Everson et al. 2016
 Tenrecidae  
  Tenrecinae  


 Echinops


   

 Setifer



   

 Hemicentetes


   

 Tenrec




   
  Geogalinae  

 Geogals


  Oryzorictinae  

 Oryzorictes 


   

 Nesogale


   

 Microgale






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The Tenrek family is now divided into three subfamilies. The Igeltenreks (Tenrecinae) form the sister group of the other Tenreks ( Reis- and Erdtenreks or Oryzorictinae and Geogalinae). The former are distinguished by their prickly, hedgehog- like fur and their short stubby tail from the latter two, which in turn are characterized by a soft-haired fur, a shrew-like exterior and a long tail. According to molecular genetic studies, the separation of the two lines took place in the Upper Eocene about 35.6 million years ago. The externally similar members of the Oryzorictinae and the Geogalinae show deviations in the dentition structure, since the Erdtenrek has a significantly reduced dentition compared to the Reistenreks. The two subfamilies split off from each other in the Lower Oligocene around 30.1 million years ago.

Sometimes the otter shrews were seen as the fourth subfamily of the Tenreks. However, a molecular genetic study from 2016 classifies them again as an independent family, which forms the sister group of the Tenreks. Both families share numerous characteristics, first and foremost the zalambdodonte bit and the missing zygomatic arch . However, there are also important differences. Unlike the tenreks, the otter shrews do not have a collarbone , the second and third toes of the rear foot are fused together and the canine tooth has two roots. The separation of the two families took place in the Lower Eocene around 48 million years ago.

One debate was whether the Malagasy tenreks form a monophyletic unit. For example, anatomical studies by Robert J. Asher from 1999 revealed a polyphyletic origin of the Malagasy tenreks in relation to the position of the otter shrews and the golden mole. Depending on the weighting of the characteristics, there was on the one hand a stronger unity of the otter shrews with the water tenrek as well as the other soft-haired tenreks and the golden mole with the hedgehog tenreks, but on the other hand there was also a deep embedding of the otter shrews in the tenreks while retaining the close relationship of the former to the water trek. Molecular genetic analyzes published shortly afterwards contradicted this and suggested that the tenreks form a clade with the otter shrews and the golden mole act as a sister group (at that time the analyzes only included a limited number of tenre cards). Subsequent genetic studies including other species confirmed this picture and led to the view that all tenreks occurring today in Madagascar go back to a common ancestor.

Overview of the recent and fossil genera of the Tenreks

The recent representatives of the Tenrek family are structured as follows:

  • Family Tenrecidae Gray , 1821 (Tenreks)
  • Subfamily Tenrecinae Gray , 1821 ( hedgehog or hedgehog )
  • Tenrec Lacépède , 1799 (Large Tenrek, 1 kind)
  • Hemicentetes Mivart , 1871 (striped duck trek , 2 species)
  • Setifer Froriep , 1806 (Large hedgehog trek, 1 species)
  • Echinops Martin , 1838 (Kleiner Igeltenrek, 1 species)
  • Oryzorictes A. Grandidier , 1870 (rice burrower, 2 species)
  • Microgale Thomas , 1882 (Kleintenreks, 21 species)
  • Nesogale Thomas , 1918 (2 types),
  • Geogale Milne-Edwards & A. Grandidier , 1872 (Erdtenrek, 1 species)

Originally, with Limnogale , another genus belonged to the Reistenreks, which contained the water tenrek ( Microgale mergulus ) and was monotypical . Molecular genetic studies of the Tenreks already carried out at the beginning of the 21st century suggested a close connection to the Lesser Tenreks, and some studies also indicated that the aquatic tenreks were deeply embedded in the genus Microgale . The latter was confirmed by a recent genetic study from 2016 that took into account all known representatives of the tenreks, so that the water tenrek was incorporated into the small tenreks.

Fossil the following forms can still be distinguished:

In 1987 the genus Ndamathaia was described on the basis of individual remains of the lower jaw from the Lower Miocene by Songhor in western Kenya , which the authors classified as a gigantic tenrek. A phylogenetic study from the year 2000, however, comes to the conclusion that these are the remains of a mongoose belonging to the extinct genus Kelba .

Research history

discovery

Panel from Flacourt's travelogue from 1658 with the first depiction of a tenrek (lower right panel, misspelled as "tendiac")
Etienne de Flacourt
Representation of the “Tanrec” (left) and the “Tendrac” (right) according to Brisson 1764

The first references to the tenreks in Western science go back to the mid-17th century and come from Étienne de Flacourt (1607–1660). The naturalist and former governor of Madagascar published his travelogue about the island (plus his stay in continental Africa) in 1658 with the work Histoire de la grande isle Madagascar , in which he also dealt with natural history. In the book, which was published several times, Flacourt described a hedgehog-like animal that would sleep for six months and dig itself into the ground. In addition, it would not eat any food during this time. Flacourt called the animal "Tendrac", but gave "Sora" as another local name. He also added a drawing that is very crude from today's perspective. It then took a good century before Georges-Louis Leclerc de Buffon (1707–1788) dealt with the Malagasy animals. Buffon published his very extensive work Histoire naturelle, générale et particulière , which includes numerous volumes , since 1749 . In the twelfth volume from 1764 he described the "Tendrac" more precisely. He also put the "Tanrec" at his side, which Buffon claims is larger than the "Tendrac". Buffon illustrated the animals with two attached drawings, using individuals from the Cabinet du roi , the predecessor of the Muséum national d'histoire naturelle in Paris , as a template and for the descriptions . These had entered the collection of the royal cabinet through a dealer from Lyon. About 13 years later, Johann Christian von Schreber (1739–1810) published his work Die Säugthiere in pictures from nature with descriptions . In this, he introduced the “Tendrac” and the “Tanrec” in German, and he also added two images of the animals, the execution of which is strongly based on the drawings from Buffon's original. In addition, Schreber awarded the scientific species names Erinaceus setosus for the "Tendrac" and Erinaceus ecaudatus for the "Tanrec". As a result, the "Tendrac" is connected to the Great Igeltenrek (today Setifer setosus ) and the "Tanrec" with the Great Tenrek (today Tenrec ecaudatus ). Buffon had depicted two striped tenreks in two additional volumes of his Histoire naturelle , which were published in 1776 and 1789, and addressed them with le jeune tanrec and le Tandrac, respectively , but these are now regarded as individuals of the actual striped tenrek and the black- headed trek .

From the Malagasy hedgehogs to the tenreks

The external appearance of the tenrek representatives known up to the end of the 18th century - it was exclusively those of the hedgehog tenreks - tempted almost all researchers of that time to interpret the animals as hedgehogs and assign them to the hedgehogs. Through Schreber's establishment of the scientific names for the Great Tenrek and the Great Igeltenrek, these were directly assigned to the genus Erinaceus and thus to the European hedgehogs (more precisely to today's small ear hedgehogs). A formal separation of the Tenreks from the genus Erinaceus was first carried out by Georges Cuvier in 1798, but he did not give a valid genus name. This is only granted to Bernard Germain Lacépède (1756–1825) a year later, who gives the name Tenrec in a taxonomic overview , other generic names should follow later. Nevertheless, the Tenreks initially remained within the group of hedgehogs, for example with Étienne Geoffroy Saint-Hilaire in 1803.

Tenrecidae versus Centetidae

John Edward Gray

The scientific genus name Tenrec , named by Lacépède in 1799, forms the basis for the name of the Tenrecidae family that is valid today. This in turn comes from John Edward Gray (1800–1875) in 1821. Gray formulated the brief description of the family as follows: Teeth, grinders, acutely tubercular; cutting, small, equal, six above, four below, canine long; body spinous; tail none. ("Teeth, molars, strictly tubercular; cutting, small, equal in size, six above, four below; canine long, body prickly; no tail."). Both the name Tenrec for the genus and the name Tenrecidae for the family were rarely used during the 19th century. Already 12 years after Lacépède, Johann Karl Wilhelm Illiger established the generic name Centetes for the great tenrek. This comes from the Greek language ( κεντητήϛ kentetes ) and is derived from κεντητος ( kentetos ) for “embroidered” or κεντεω ( kenteo ) for “sting”. Centetes was the godfather for the family name of the Centetidae. Andrew Murray introduced this in 1866, in which he combined all of the Tenre cards known at the time (Murray named Centetes , Ericulus and Echinogale ; these are now listed under Tenrec , Setifer and Echinops ). Murray's scientific name for the family was preceded by the term Centetina, coined by Charles Lucien Jules Laurent Bonaparte in 1838. Bonaparte led Centetina as a subgroup within the hedgehogs, but thereby separated the tenreks from the actual hedgehogs. The term Centetidae remained widespread through the remainder of the 19th century and was also used in the first half of the 20th century. In 1931 and 1945, George Gaylord Simpson pointed out in his classification schemes of mammals that Tenrec is the older genus name compared to Centetes , Tenrecidae was also named before Centetidae. As a result, the two names assigned later are invalid and should only be regarded as synonyms . Nevertheless, Centetes and Centetidae remained in use until the 1960s.

The tenreks as insect eater

The close relationship between the hedgehogs and the tenreks, which was assumed in the late 18th and early 19th centuries, led to the fact that they were also merged in the superordinate system. In his work from 1811 Illiger had set up the group of the Subterranea ("subterranean") and in it united both the hedgehogs and tenreks as well as the golden mole , the moles and the shrews , he created a taxon that is no longer recognized today that has all the forms united, which would later be led under the term "insect eater". The insectivores ( Les Insectivores ) were first introduced informally by Cuvier in 1817. He divided them into two groups: one with long internal incisors, in which he placed the golden mole, shrews and hedgehogs, and a second with large canine teeth, for example in the moles and the tenreks. The concept was adopted four years later by Thomas Edward Bowdich with the introduction of the scientific name Insectivora, which resulted in the recognition of the insectivore as a classification group.

During the 19th and much of the 20th centuries, the concept of the insect eater was hardly questioned. New findings formed the basis for an ever finer division of the group. Wilhelm Peters worked out two nameless groups in 1863, the much larger one struck by the lack of an appendix , while a smaller one had such an organ. The latter includes the elephants and the shrews , the former the rest of the insectivora including the tenreks and the golden mole. Later, in the second volume of his work General Morphology of Organisms , Ernst Haeckel gave the two groups the official names Lipotyphla (without appendix) and Menotyphla (with appendix). Theodore Gill took a completely different approach in the 1880s. He differentiated the insectivores according to the expression of the chewing pattern of the molars. Accordingly, there were two types, on the one hand the zalambdodonta, which have a V-shaped enamel ridge on the molars (from the Greek ζα ( za- ) for an excess and the Greek letter Λ (lambda)), on the other hand the dilambdodonta with a W-shaped ridge ( of Greek δι ( di "two") and the Greek letter Λ (lambda)). According to Gill, the tenreks belonged to the zalambdodont insectivores, a position they shared with the gold mole and the slit weevil. All other insectivores formed the dilambdodonte group. With this, Gill recognized for the first time a closer relationship between the Tenreks and the Goldmullen.

William K. Gregory made a further attempt at structuring in 1910 in his work The orders of Mammals , in doing so he relied in part on the work of Haeckel and Gill and combined them. So Gregory separated the menotyphla from the insectivores, he led the rest of the group under the name Lipotyphla. He subdivided these in turn and kept the tenreks and gold moles within the Zalambdodonta to Gill, while he divided the remaining Dilambdodonta into the two large groups Erinaceomorpha (hedgehogs) and Soricomorpha (shrews and moles). In his general taxonomy of mammals published in 1945, George Gaylord Simpson subdivided the insectivores into five recent superfamilies, the tenreks forming the tenrecoidea, the gold mole the chrysochloroidea, he placed both in relative proximity to one another. The closer relationship between the golden mole and the Tenreks was also often discussed later, for example by Percy M. Butler in 1956. Butler partially adopted the systematic scheme of insectivores from Gregory, but he only differentiated the two large groups of the Erinaceomorpha and the Soricomorpha, the Tenreks and the gold mole he assigned to the Soricomorpha. In the 1960s, Leigh Van Valen separated the zalambdodonta, i.e. the tenreks, golden mole and weevil, from the insectivores and placed them in the order of the Deltatheridia that he had created. Van Valen grouped the Zalambdodonta opposite the so-called Hyaenodonta. The latter consisted of several extinct families, including the Hyaenodontidae and the Oxyaenidae ; In a more classic view, these belong to the non-self-contained group of the " Creodonta " ("primal carnivores"). The view was largely not adopted. Subsequent authors such as Malcolm C. McKenna in the 1970s and Michael J. Novacek in the 1980s kept the zalambdodont forms mostly with the insectivores and there with the Soricomorpha. With regard to the Tenreks McKenna repeated this view together with Susan K. Bell again in his overview work on the systematics of mammals in the 1990s, but here he excluded the gold mole from the Soricomorpha and placed it in its own order.

New insights

It was not until the transition from the 20th to the 21st century that new knowledge gained acceptance with the development of new research methods. In 1998, a team and Michael J. Stanhope published molecular genetic studies on insectivores for the first time . It turned out that the insectivores do not form a self-contained unit. Accordingly, the moles, hedgehogs and shrews represent a common group, which have a closer relationship to the pair and odd-toed ungulates as well as to the pancake , predatory and bats (the group was later called Laurasiatheria ). The tenreks, on the other hand, form a unit with the gold mole, which was already known for anatomical reasons, and also with the elephants, the aardvark, the elephants, the snakes and the manatees. Since these are mainly groups of animals that are endemic to Africa, Stanhope and colleagues named them Afrotheria. At the same time, they relegated the African branch of the original insectivores, consisting of the tenreks and golden mole, to an independent order within the Afrotheria, the tenre-kitties (Afrosoricida; actually translated "African shrews"). Three years later, the tenre-kitties, the elephants and the aardvark were combined in a higher-level taxon called Afroinsectiphilia . In the period that followed, further analyzes supported the result, including the isolation of a specific retroposon , the so-called Afro SINE , which all representatives of Afrotheria have in common. Accordingly, the similarities of the tenreks and golden mole to other representatives of the insectivores are only to be understood as a convergence .

Tribal history

On the origin of the Malagasy tenreks

The oldest fossil finds of the Tenreks come from the Eocene in Africa, from Madagascar only subfossil remains from the Holocene are available. The origin of the Tenreks is therefore most likely to be found in Africa. Madagascar separated from Africa around 160 million years ago and has existed as an independent island for at least 80 to 90 million years. Today it is separated from the continent by the approximately 400 km wide road from Mozambique . According to the molecular genetic data, all Malagasy tenreks go back to an original form. According to the molecular clock , the family originated around 48.3 million years ago, and the tenreks began to fanned out around 35.6 million years ago. Between these two corner points, Madagascar can be assumed to have been colonized by early forms of the Tenreks. How this happened is unclear, but the possibility of a temporary land bridge through the Mozambique Strait, possibly along the Davie Fracture Zone , a fault zone off the island's western coast, has been suggested. The land bridge may have existed from the Eocene to the Lower Miocene as a result of increased seismic activity . Another possibility would be to cross the water surface on driftwood.

The number of Tenreks immigrating to Madagascar is unclear. Originally, several waves of colonization were assumed, which is largely due to the unclear relationships between the Tenreks in the late 20th century. From today's perspective, the recent Tenreks are monophyletic, which means that a single settlement would be sufficient. However, the fossil forms from the early Miocene of Africa, which from an anatomical point of view show a closer relationship to the Erdtenrek, are problematic. If so, the biogeographical development history of the Tenreks is more complicated than just presented. Then at least a second migratory movement (from the mainland to Madagascar or vice versa) may have to be assumed. However, this question cannot yet be answered.

After the colonization of Madagascar, there was rapid diversification. Based on today's Tenreks, the speciation rate in the humid tropical forests of the eastern part of the island was particularly high. Adaptations to the drier conditions in the west were made several times independently, both in the group of the hedgehog treks and among the soft-haired forms. In the course of the colonization of Madagascar by the Tenreks, there was apparently a reduction or a “blurring” of gender-specific size characteristics, which are sometimes more pronounced among continental representatives of Afrotheria. The reasons for this - whether they can be traced back to a specific founder population ( founder effect ) or to adaptations to the conditions of an isolated land mass ( ecologically controlled evolutionary development ) - have not yet been clarified. A similar phenomenon occurs with the lemurs of Madagascar.

Eocene

The oldest known tenrek fossils to date came to light in the northern restricted area in Namibia . There are numerous skull and jaw remains as well as parts of the musculoskeletal system from the Eocliff site , a travertine deposit that had formed in the Upper Eocene at the point where a calcareous spring emerged. The findings represent two genera, one hand Sperrgale other hand Arenagale . Both forms are small representatives of the tenreks, roughly the size of today's earth tenreks. They are characterized by a more primitive dentition, in which the zalambdodontal chewing surface pattern is not yet quite as distinct due to the metaconus of the maxillary molars, which is clearly separated from the paraconus. The musculoskeletal system suggests that the limbs could be used very mobile, so there is no evidence of a digging way of life. A lower jaw branch from Nanogale is also available from the restricted area . It came to light in the limestone layers of the Black Crow site , which are somewhat older than the Eocliff . Morphologically, through the design of the ascending branch on the lower jaw , Nanogale can not be directly connected to the Tenreks, but is part of the wider relationship environment, the Tenrecomorpha. With a length of only 8.6 mm, the lower jaw is comparable to that of the smallest known mammals that do not belong to the bats , such as various representatives of the thick-tailed shrews .

Oligocene

In the further course of the Eocene and Lower Oligocene , no direct members of the Tenreks are found, but some closely related forms can be proven. The Fayyum fossil deposit in northern Egypt is of importance here . Dilambdogale , which was described in 2010, comes from the Qasr-el-Sagha formation and thus from the uppermost section of the Eocene . Other forms were found with Widanelfarasia , Jawharia and Qatranilestes from the Gebel-Qatrani formation and belong to the Oligocene. All four representatives are largely only known from mandibular and maxillary fragments, in very few cases there is also postcranial skeletal material. These relatively primitive representatives also had a separate para and metaconus on the upper molars, and a W-shaped enamel ridge pattern was also formed on the foremost molar, which is reminiscent of the dilambdodontic chewing surface structure of insectivores. The posterior molars, on the other hand, had a zalambdodontic pattern, with which the shapes assume a transition position to the typical Afrosoricida.

Miocene

Only in the following Miocene are Tenreks clearly documented again. Most of the fossil remains discovered so far come from East Africa . They have been found in the Legetet Formation , the Hiwegi Formation and the Kapurtay Agglomerates , which are around 18 to 20 million years old. A total of three genera have been handed down in the individual exposures. Parageogals , documented over a partial skull and several dentition fragments, possibly represents a close relative of the Erdtenreks . It differs from the recent form by its slightly larger dimensions and its even more reduced dentition. In the skull structure, however, it shows some primeval features. Erythrozootes was described on the basis of a partial skull; several teeth and jaw fragments are also known. The shape was larger than the Kleintenreks . The most striking features are the very large front upper incisors, a very well developed protoconus on the maxillary molars and a slightly reduced talonid on the molars of the lower jaw. In these features true Erythrozootes with the third fossil genus, Protenrec , agreed, but the latter was much smaller and had next to a larger bite several different skull features, such as the relatively long channel of the infra-orbital foramen or very far forward shifted lacrimal foramen . From Protenrec far more mandibular remains and a front head part have been documented outstanding, the reference is Songhor in western Kenya . In addition, individual parts of the lower jaw and skull of the genus came to light in Arrisdrift as well as in Elisabethfeld and Langental in the restricted area of ​​Namibia, the animals were on average slightly smaller than those from East Africa. The sites date to the Lower and the beginning of the Middle Miocene. Unlike Parageogale , relationships continue to be Erythrozootes and Protenrec more difficult to determine, they are sometimes placed in its own subfamily of Protenrecinae, but perhaps is also in close relationship with the large-eared tenrec. From the Lower Miocene, Elisabethfeld discovered a previously single lower jaw of Promicrogale in the same region ; the genus shows relationships with the Kleintenreks.

Plio and Pleistocene

Neither the Pliocene nor the Pleistocene have been found to date from Tenreks in Africa or Madagascar.

Holocene

The earliest evidence of fossil records of the Tenreks in Madagascar dates back to the Holocene , with most of the finds being less than 2000 years old. Bones from members of the family were found in several cave sites, the most important of which are the Anjohibe Cave in the northwest, the Ankilitelo Cave in the southwest and the Andrahomana Cave in the southeastern part of the island. All three are characterized by an extraordinary wealth of fossils in mammals, such as bats , rodents , predators and primates . The tenreks are represented with around a dozen species, which can be referred to the hedgehog treks as well as to the terrestrial treks and to the rice treks. These include some species such as the major long-tailed Kleintenrek or the Nasolo-Kleintenrek , whose current range is significantly limited so that they no longer occur in the corresponding find region. In addition, with Microgale macpheei an extinct relatives could be proven, he has so far only been documented from the Andrahomana cave. In addition to the actual Tenreks, the closely related form Plesiorycteropus with at least two species appeared in the course of the Holocene . It is known from at least a dozen sites, which are mainly distributed in central and western Madagascar. The extinction of individual members of the Tenreks or closely related forms, but also the withdrawal of some species from the former range, has various reasons. These are related on the one hand to the increasing drying up of the western part of the island in the last two to three thousand years, and on the other hand to the arrival of humans during this period.

Threat and protection

The greatest threat to the existence of the individual tenre cards as well as to many other forest-dwelling animal species in Madagascar is the ongoing destruction of their habitat. The IUCN lists six species as "endangered" ( endangered ) or "endangered" ( vulnerable ), these are exclusively representatives of the small tenreks with a clearly limited range. Another species from the genus of the rice burrower is classified in the category " data deficient " due to a lack of information . The Igeltenreks, on the other hand, are widespread and have become used to being close to humans as cultural followers . They are therefore currently not considered to be threatened. However, due to their size, they are more exposed to human hunting. Their meat is mostly used as a food resource. Almost all tenre cards are found in nature reserves.

literature

  • JF Eisenberg and Edwin Gould: The Tenrecs: A Study in Mammalian Behavior and Evolution. Smithsonian Institution Press, 1970, pp. 1-138
  • Nick Garbutt: Mammals of Madagascar. A complete guide. Yale University Press, 2007, pp. 1–304 (pp. 32–56)
  • Konrad Herter: About the hedgehogs of Madagascar (Tenrecinae). Meeting reports of the Society of Friends of Natural Sciences in Berlin NF 2, 1962, pp. 5-37
  • Paulina D. Jenkins: Tenrecidae (Tenrecs and Shrew tenrecs). 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. 134–172 ISBN 978-84-16728-08-4
  • Ronald M. Nowak: Walker's Mammals of the World . The Johns Hopkins University Press, Baltimore 1999, ISBN 0-8018-5789-9 .
  • Link E. Olson: Tenrecs. Current Biology 23 (1), 2013, pp. R5-R8

Individual evidence

  1. ^ A b Peter M. Kappeler, Charles L. Nunn, Alexander Q. Vining and Steven M. Goodman: Evolutionary dynamics of sexual size dimorphism in nonvolant mammals following their independent colonization of Madagascar. Scientific Reports 9, 2019, p. 1454 doi: 10.1038 / s41598-018-36246-x
  2. ↑ a b c d e f Nick Garbutt: Mammals of Madagascar. A complete guide. Yale University Press, 2007, pp. 1–304 (pp. 32–56)
  3. ↑ a b c d Konrad Herter: About the hedgehogs of Madagascar (Tenrecinae). Meeting reports of the Society of Friends of Natural Sciences in Berlin NF 2, 1962, pp. 5-37
  4. ↑ a b c d e f g h i j k J. F. Eisenberg and Edwin Gould: The Tenrecs: A Study in Mammalian Behavior and Evolution. Smithsonian Institution Press, 1970, pp. 1-138
  5. ↑ Sive Finlay and Natalie Cooper: Morphological diversity in tenrecs (Afrosoricida, Tenrecidae): comparing tenrec skull diversity to their closest relatives. PeerJ 3, 2015, p. E927 doi: 10.7717 / peerj.927
  6. ↑ a b c Wilhelm Leche: On the history of the development of the mammalian tooth system, at the same time a contribution to the tribal history of this group of animals. Part two: phylogeny. Second booklet: Families of the Centetidae, Solenodontidae and Chrysochloridae. Zoologica 20, 1906/1908, pp. 1–157 ( [1] )
  7. ^ A b Steven M. Goodman and Paulina D. Jenkins: The Insectivores of the Réserve Spéciale d'Anjanaharibe-Sud, Madagascar. Fieldiana Zoology 90, 1998, pp. 139-161
  8. ^ A b c Percy M. Butler: Macroscelidea, Insectivora and Chiroptera from the Miocene of East Africa. Palaeovertebrata 14 (3), 1984, pp. 117-200
  9. ^ Robert J. Asher and Marcelo R. Sánchez-Villagra: Locking Yourself Out: Diversity Among Dentally Zalambdodont Therian Mammals. Journal of Mammalian Evolution. 12 (1/2), 2005, pp. 265-282
  10. ↑ a b Link E. Olson: Tenrecs. Current Biology 23 (1), 2013, pp. R5-R8
  11. ^ Justine A. Salton and Frederick S. Szalay: The Tarsal Complex of Afro-Malagasy Tenrecoidea: A Search for Phylogenetically Meaningful Characters. Journal of Mammalian Evolution 11 (2), 2004, pp. 73-104
  12. ^ Justine A. Salton and Eric J. Sargis: Evolutionary morphology of the Tenrecoidea (Mammalia) carpal complex. Biological Journal of the Linnean Society, 93, 2008, pp. 267-288
  13. ^ Justine A. Salton and Eric J. Sargis: Evolutionary Morphology of the Tenrecoidea (Mammalia) Forelimb Skeleton. In: EJ Sargis and M. Dagosto (Eds.): Mammalian Evolutionary Morphology: A Tribute to Frederick S. Szalay, Springer Science, 2008, pp. 51-71
  14. ^ Justine A. Salton and Eric J. Sargis: Evolutionary Morphology of the Tenrecoidea (Mammalia) Hindlimb Skeleton. Journal of Morphology 270, 2009, pp. 367-387
  15. ^ Howard P. Whidden: Extrinsic Snout Musculature in Afrotheria and Lipotyphla. Journal of Mammalian Evolution. 9 (1/2), 2002, pp. 161-184
  16. ↑ Hideko Endo, Takahiro Yonezawa, Felix Rakotondraparany, Motoki Sasaki and Masami Hasegawa: The adaptional strategies of the hindlimb muscles in the Tenrecidae species including the aquatic web-footed tenrec (Limnogale mergulus). Annals of Anatomy 188, 2006, pp. 383-390
  17. ↑ H. Endo, M. Oishi2, T. Yonezawa, F. Rakotondraparany and M. Hasegawa: The Semifossorial Function of the Forelimb in the Common Rice Tenrec (Oryzorictes hova) and the Streaked Tenrec (Hemicentetes hemispinosus). Anatomie, Histology, Embryology 36, 2007, pp. 413-418
  18. ↑ B. Riedelsheimer, Pia Unterberger, H. Künzle and U. Welsch: Histological study of the cloacal region and associated structures in the hedgehog tenrec Echinops telfairi. Mammalian Biology 72, 2007, pp. 330-341
  19. ^ Peter Langer: The digestive tract and life history of small mammals. Mammal Review 32 (2), 2002, pp. 107-131
  20. ↑ a b c J. J. Petter and A. Petter-Rousseaux: Notes biologiques sur les Centetinae. La Terre et la Vie 17 (1), 1963, pp. 66-80
  21. ↑ J. Michael Bedford, Orin B. Mock and Steven M. Goodman: Novelties of conception in insectivorous mammals (Lipotyphla), particularly shrews. Biological Review 79, 2004, pp. 1-19
  22. ↑ AC Enders, TN Blankenship, SM Goodman, V. Soarimalala and AM Carter: Placental Diversity in Malagasy Tenrecs: Placentation in Shrew Tenrecs (Microgale spp.), The Mole-Like Rice Tenrec (Oryzorictes hova) and The Web-Footed Tenrec ( Limnogale mergulus). Placenta 28, 2007, pp. 748-759
  23. ↑ AM Carter, TN Blankenship, H.Künzle and AC Enders: Structure of the Definitive Placenta of the Tenrec, Echinops telfairi. Placenta 25, 2004, pp. 218-232
  24. ↑ Jason A. Kaufman, Gregory H. Turner, Patricia A. Holroyd, Francesco Rovero and Ari Grossman: Brain Volume of the Newly-Discovered Species Rhynchocyon udzungwensis (Mammalia: Afrotheria: Macroscelidea): Implications for Encephalization in Sengis. PlosOne 8 (3), 2013, p. E58667 doi: 10.1371 / journal.pone.0058667
  25. ↑ Leah Krubitzer, Heinz Künzle and Jon Kaas: Organization of Sensory Cortex in a Madagascan Insectivore, the Tenrec (Echinops telfairi). Journal of Comparative Neurology 379, 1997, pp. 399-414
  26. ^ Susanne Radtke-Schuller and Heinz Künzle: Olfactory bulb and retrobulbar regions in the hedgehog tenrec: Organizations and interconnections. Journal of Comparative Neurology 423 (4), 2000, pp. 687-705.
  27. ↑ Heinz Künzle: thalamo-striatal Projections in the hedgehog tenrec. Brain Research 1100, 2006, pp. 78-92
  28. ↑ a b c d e f g h i j k Kathryn M. Everson, Voahangy Soarimalala, Steven M. Goodman and Link E. Olson: Multiple loci and complete taxonomic sampling resolve the phylogeny and biogeographic history of tenrecs (Mammalia: Tenrecidae) and reveal higher speciation rates in Madagascar's humid forests. Systematic Biology 65 (5), 2016, pp. 890-909 doi: 10.1093 / sysbio / syw034
  29. ↑ Claudette Patricia Maminirina, Steven M. Goodman and Christopher J. Raxworthy: Les microammifères (Mammalia, Rodentia, Afrosoricida et Soricomorpha) du du massif Tsaratanana et biogeography of Forests de montagne de Madagascar. Zoosystema 30 (3), 2008, pp. 695-721
  30. ↑ a b Melanie Dammhahn, Voahangy Soarimalala and Steven M. Goodman: Trophic Niche Differentiation and Microhabitat Utilization in a Species-rich Montane Forest Small Mammal Community of Eastern Madagascar. Biotropica 45 (1), 2013, pp. 111-118
  31. ^ RDE MacPhee: The Shrew Tenrecs of Madagascar: Systematic Revision and Holocene Distribution of Microgale (Tenrecidae, Insectivora). American Museum Novitates 2889, 1987, pp. 1-45
  32. ^ A b Edwin Gould: Evidence for echolocation in the Tenrecidae of Madagascar. Proceedings of the American Philosophical Society 109 (6), 1965, pp. 352-360
  33. ↑ Olga V. Zherebtsova: Morphofunctional interpretation of the quills stridulating in tenrecs (Lipotyphla, Tenrecidae). Russian Journal of Theriology 5 (1), 2006, pp. 1-11
  34. ↑ Danielle L. Levesque, D. Rakotondravony and Barry G. Lovegrove: Home range and shelter site selection in the greater hedgehog tenrec in the dry deciduous forest of Western Madagascar. Journal of Zoology 287, 2012, pp. 161-168
  35. ↑ Walter Poduschka: Augendrüsensekretion in Tenreciden Setifer setosus (Froriep 1806), Echinops telfairi (Martin 1838); Microgale dobsoni (Thomas 1918) and M. talazaci (Thomas 1918). Zeitschrift für Tierpsychologie 35, 1974, pp. 303-319
  36. ^ A b Edwin Gould and John F. Eisenberg: Notes on the biology of the Tenrecidae. Journal of Mammalogy 47 (4), 1966, pp. 660-686
  37. ↑ Jonathan Benstead, Kevin H. Barnes and Catherine M. Pringle: Diet, acvtivity patterns, foraging movements and response to deforestation of the aquatic tenrec Limnogale mergulus (Lipotyphla: Tenrecidae) in eastern Madagascar. Journal of Zoology 254, 2001, pp. 119-129
  38. ^ A b c Peter J. Stephenson and Paul A. Racey: Reproductive energetics of the Tenrecidae (Mammalia: Insectivora). I. Large-eared tenrec, Geogale aurita. Physiological Zoology 66 (5), 1993, pp. 643-663
  39. ^ A b Peter J. Stephenson and Paul A. Racey: Reproductive energetics of the Tenrecidae (Mammalia: Insectivora). II. The shrew-tenrecs, Microgale spp. Physiological Zoology 66 (5), 1993, pp. 664-685
  40. ^ PJ Stephenson, JR Speakman and PA Racey: Field metabolic rate in two species of shrew-tenrec, Microgale dobsoni and Microgale talazaci. Comparative Biochemistry and Physiology 107A (2), 1994, pp. 283-287
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