Erdtenrek

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Erdtenrek
Systematics
without rank: Afroinsectiphilia
Order : Tenrecus (Afrosoricida)
Family : Tenreks (Tenrecidae)
Subfamily : Geogalinae
Genre : Geogals
Type : Erdtenrek
Scientific name of the  subfamily
Geogalinae
Trouessart , 1879
Scientific name of the  genus
Geogals
Milne-Edwards & A. Grandidier , 1872
Scientific name of the  species
Geogale aurita
Milne-Edwards & A. Grandidier , 1872

The large-eared tenrec ( Geogale aurita ), also Erdtanrek , Zwergtanrek or large ear Tenrek called, is a mammal of the family of tenrecs (Tenrecidae). It occurs in western and southern Madagascar and inhabits dry forests and open landscapes. Overall, however, it is rather rare. The animals are among the smallest tenreks of all. Outwardly they resemble the Kleintenreks , their most noticeable feature are the large, mouse-like ears. The fur has various light brown shades, the tail is only about half the length of the rest of the body.

The way of life is relatively well researched in some aspects. The animals live on the ground and are nocturnal. They are considered extremely cold-blooded, the body temperature adapts to the outside temperature, and this is associated with a low metabolic rate . Especially in the dry season, there is a daily cold rigidity ( torpor ). The diet consists of invertebrates such as insects . The length of the gestation period is variable and is influenced by the low metabolism and the outside temperature. A litter comprises one to five newborns who are born as nestlings .

The Erdtenrek is not an endangered species. The species was described in 1872. Especially in the course of the 20th century it was often assigned to the Reistenreks , today it is mostly in its own subfamily (Geogalinae). Several subfossil finds from Madagascar are known, and some fossil remains have been found in East Africa that are more closely related to the Erdtenrek and date to the Lower Miocene .

features

Habitus

The Erdtenrek is one of the smallest representatives of the Tenreks . It reaches a total length of 9.0 to 10.7 cm, the head-trunk length is 6.1 to 7.6 cm, the tail length is 3.4 to 4.1 cm. The body weight varies from 5.0 to 8.5 g. In general, the animals resemble the Kleintenreks ( Microgale ) in their external appearance . The body is spindle-shaped and the snout pointed in front. The most noticeable hallmarks are the large, mouse-like ears that are 15 to 18 mm long. They have a brownish color and are covered on both sides by a very short and fine, mostly gray fur. The very long vibrissae extend diagonally backwards to the top, their basic color is brown. They appear more numerous on the snout than on the lower jaw. The body fur is soft, short and dense. The color of the top ranges from light brown to light reddish brown. The underside is mostly tinted creamy white. Orange spots are sometimes formed on the sides. The tail is scaly and covered with fine hair. The limbs end in front and back in five rays each, which are delicately built and have small claws. The hind foot is relatively small in relation to the length of the head and torso, its length is only 11 to 12 mm. Females have one or two pairs of teats in the chest, a maximum of one pair in the abdomen and two pairs in the groin.

Skull and dentition features

The skull becomes 19 to 20 mm long and on the cranium 7.5 to 8.0 mm wide. Overall, it appears elongated, the rostrum is conspicuously drawn in shortly before the molars and is narrow, the posterior area of ​​the skull appears short and angular. In the side view, the forehead line is straight and rising evenly from front to back. The area behind the eyes is narrow and elongated. As with all tenreks, a closed zygomatic arch is missing , but the anterior arch attachment is relatively wide. Strong muscle attachment marks are formed on the occiput. The dentition consists of a total of 34 teeth, which differs from the Kleintenreks and the rice rooters ( Oryzorictes ). The dental formula is: . The first incisor in each case in the upper and lower row of teeth clearly protrudes over the following and the canine . In the upper dentition, the first and second incisors are separated by a short diastema . There are additional cusps on the crowns of the front teeth. The two anterior upper premolars (P2 and P3) and the first lower premolar are very small, while the lower posterior premolars have a distinct protoconid. The molars have a zalambdodontic occlusal surface pattern, which consists of three main cusps (para-, proto- and metaconus; related to the maxillary molars) in a triangular arrangement. In the upper jaw, the first two molars show noticeable narrowing anterior and posterior. These are also evident in the last one, but this is also stretched to the side. The third lower molar has only a small talonid (a deep protrusion on the tooth surface).

Skeletal features

The skeleton structure of the earth tenreks is very generalized for tenreks and shows hardly any special features. In the hand in particular, there are hardly any characteristics that point to a certain specialization in the way of moving and living. A deviation can be found on the shoulder blade , where the metacromion, an extension of the acromion, is not oriented forwards, but backwards and thus parallel to the longitudinal axis of the shoulder blade. The construction of the hind legs mediates between the hedgehog treks (Tenrecinae) and the rice treks (Oryzorictinae). Like the former, the Erdtenrek has an elongated pelvis that is almost the length of the thigh bone , the femoral head is also large and the neck rather short, and the third rolling mound is small. On the other hand, the high-seated small rolling mound on the femur and the extraordinarily slim or long shin rather refer to the trek. A comparatively similar mixture of characteristics is also pronounced on the foot skeleton, for example on the ankle and heel bones as well as on the ankles.

distribution

Distribution area of ​​the Erdtenreks

The Erdtenrek is endemic in Madagascar, it occurs there mainly in the southern and western parts of the country. The number of sites with evidence of the Erdtenreks is limited, however, field investigations since the 1990s have found it in a little more than a dozen places, the majority of which are in the province of Toliara . The forest area of ​​Kirindy near Morondava is an important site in the west . In the southwest, animals have been observed in the Zombitse-Vohibasia National Park , in the Tsimanampetsotsa National Park south of Toliara , in the Beza Mahafaly nature reserve and in the Mikea forest. The distribution area is limited in the southeast by the western foothills of the Anosyenne Mountains , in the west it extends northwards to the Tsiribihina river. The species was also reported from the Ankarafantsika National Park in northwest Madagascar in 2002 , but it seems that it has been confused with a young of the short-tailed lesser tenreks ( Microgale brevicaudata ). The entire west and south of Madagascar is characterized by a very dry climate, in some cases with annual rainfall of only 400 to 500 mm. The Erdtenrek inhabits dry forests , gallery forests and open thorn bush landscapes or savannah areas . In general, the species is considered rare. It is documented most frequently from Beza Mahafaly, where gallery forests consisting of tamarind trees and legumes dominate.

Way of life

Territorial behavior

More extensive data is only available in individual aspects about the way of life of the Erdtenrek. It is strictly nocturnal and lives predominantly solitary, however, males and females have often been observed close together. The animals spend the rest phases in underground burrows in the sandy subsoil, sometimes near dead tree trunks.

Diet and energy balance

When looking for food, the animals use their good sense of smell and hearing (the sense of sight is only weakly developed). The large ears constantly move back and forth in search of prey. The main diet consists of invertebrates , often grasshoppers and termites .

The Erdtenrek is one of the warmest representatives of the Tenreks, the body temperature equals the outside temperature in every season. With ambient temperatures varying from 18 to 27 ° C, the body temperature is 20.8 to 33.2 ° C. In the upper area, the body temperature is around 4 ° C below that of the Kleintenreks . There are no differences between male and female animals, nor between pregnant or lactating females and those outside the reproductive phase, only individual individuals sometimes show slightly higher body temperatures during the carrying of the offspring. Associated with this is a low metabolic rate , which only reaches 53% of the value that would be expected for mammals of roughly the same size. There are no major fluctuations over the course of the day or year. This differs from the Kleintenreks, which show an increased metabolism at least under physical stress, for example during the reproductive phase. The extremely low metabolism of the Erdtenrek is possibly due to the termite-based diet and the very dry climatic conditions in western Madagascar. In unfavorable environmental conditions, such as during the dry season, a daily cold rigidity ( torpor ) occurs.

Reproduction

Reproduction is well documented based on laboratory research in the late 1980s. It takes place mostly from late September to March. A characteristic of sexual intercourse is that the male's swollen penis is held in the female's vagina after fertilization , which can last for up to 21 minutes. The gestation period of the Erdtenreks lasts from 54 to 69 days, which roughly corresponds to the other tenreks, but is extremely long due to the small size of the animals. The large variation in the length of the gestation period is caused by the outside temperature and the associated fluctuation in body temperature, as well as by the low metabolism. The torpor phases that set in at low outside temperatures cause a respective interruption in embryonic development , which was previously only known from some bats such as the pipistrelle . The ability to carry offspring with fluctuating body temperatures and a low metabolic rate also likely minimizes the overall cost of reproduction. During the gestation period, the female increases in weight considerably, starting from an average weight of 5.9 g, the body weight of the pregnant animals increased by around 4.1 g, which corresponds to about 68% more body mass.

The birth of the offspring takes place between November and March. A litter includes one to five newborns, who are born as nestlings naked and with their eyes and ears closed. The average birth weight of a newborn from a litter with three boys is 0.83 g, with a litter of five boys it is 0.63 g. Accordingly, the offspring in larger litters are on average lighter than those in smaller ones. The initial weight gain varies from 0.05 to 0.08 g, the increase in length varies from 0.7 to 1 mm. Here, too, there is a dependency on the litter size, as boys from smaller litters grow faster than those from larger litters. The eyes open after 21 to 26 days, rarely this can last up to the 33rd day. The time of weaning cannot be clearly determined as the young start to eat solid food beforehand. Usually 24 to 51 days are accepted. It is noteworthy that dams re-enter the oestrus immediately after the birth of the offspring and can be fertilized (postpartum oestrus), so that while they are still raising and nursing the offspring, they give birth to another. The advantage lies in the higher reproduction rate and the optimal use of the good environmental conditions during the reproductive phase. Life expectancy in the wild is not known; animals have lived to be two and a half years old in human care.

Predators

The most important predators are the barn owl and the Madagascar eared owl . According to investigations into the Beza Mahafaly nature reserve , the earthenrec is only a subordinate prey for both birds there. In the case of the barn owl, the proportion of individuals of the earthenrec in the entire range of prey reached around 2.8%, the biomass proportion 1.3%. For the Madagascar eared owl, the corresponding values ​​are 5.3 and 0.6%, respectively. Analyzes of the barn owl's bulges in the Tsimanampetsotsa National Park and in the Kirindy forest came to comparable results . In contrast, the barn owl's earth tenrek in the vicinity of Beahitse in the extreme south-west of Madagascar falls significantly more often, the tenrekart is represented in the dunes with an individual proportion of 15.9% and a biomass proportion of 4.3%, according to studies. Other potential predators are the narrow-striped mongoose and the Madagascar plump-nose snake .

Systematics

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


 Echinops


   

 Setifer



   

 Hemicentetes


   

 Tenrec




   
  Geogalinae  

 Geogale aurita


  Oryzorictinae  

 Oryzorictes 


   

 Nesogale


   

 Microgale






Template: Klade / Maintenance / Style

The Erdtenrek is a species from the genus Geogale , which is currently considered monotypical . Geogale, in turn, belongs to the Tenrek family (Tenrecidae). Within the Tenreks, Geogale is again the only member of the subfamily of the Geogalinae, one of the three main lines of the Tenreks. The other two are represented by the spiky hedgehog treks (Tenrecinae) and the extremely varied, soft- haired rice treks (Oryzorictinae). Due to some characteristics such as nocturnal activity, a small body size, a poor sense of sight, a low metabolic rate and low body temperature, but also the formation of abdominal mammary glands, the Erdtenrek is considered to be very primitive. According to molecular genetic analyzes, the Reistenreks form the sister group of the Erdtenreks. The representatives of both groups are outwardly very similar due to their general size and their soft coat, but a significant difference is the different set of teeth. They were separated from each other in the Lower Oligocene more than 30 million years ago.

As a rule, no subspecies of the Erdtenreks are distinguished. In 1930, however, Guillaume Grandidier and Gabriel Petit established the subspecies Geogale aurita orientalis , based on an individual from Fenoarivo in the Toamasina province . This rather unusual point of distribution of the Erdtenreks in the humid east of Madagascar could not be confirmed in field investigations on site, so that the status of the subspecies is currently considered uncertain.

The Erdtenrek was first scientifically described in 1872 by Alphonse Milne-Edwards and Alfred Grandidier . The specimens used for this had been caught from holes in the ground, three individuals came from the area around Morondava , another from Toliara , the former site is now regarded as a type locality of the species. The scientific generic name Geogale (from the Greek words γῆ ( gē̂ ) for "earth" and γαλἑη ( gale ) for "weasel") refers to the underground way of life of the animals, the species name aurita (from the Latin word auris for "ear") refers on the particularly large ears. Milne-Edwards and Grandidier did not name a holotype specimen and gave neither the size nor the sex of the individuals. It was not until almost sixty years later that Guillaume Grandidier and Gabriel Petit presented an extensive skeletal anatomical and soft tissue morphological description of the Erdtenreks. The name that is used today for the subfamily, Geogalinae, was introduced by Édouard Louis Trouessart in 1879, but the associated publication was not published until two years later. He referred the group to the otter shrews (Potamogalidae).

Research history

To the genealogical position

Trouessart's positioning of the Geogalinae and thus of the Erdtenreks within the otter shrews remained largely in place until the end of the 19th century, for example with George Edward Dobson in 1883. In 1885, Theodore Gill explained the close relationship of the Erdtenreks with the African otter shrews with the shape of the Palatine bone and the arrangement of the teeth. But he also noticed the clear similarities to the Kleintenreks, within his classification he referred the Geogalinae to the family rank (like the other subfamilies of the Tenreks). At the beginning of the 20th century, in 1907, Wilhelm Leche placed the Erdtenrek in the Tenrek family and added it to the Reistenreks, from which it hardly differs externally. The Erdtenrek largely retained this position in the further course of the 20th century, for example with George Gaylord Simpson in 1945, Leigh Van Valen in 1967 and Henri Heim de Balsac or John F. Eisenberg and Edwin Gould in the 1970s. In 2006 Robert J. Asher and Michael Hofreiter then substantiated the assignment with the help of genetic data, which suggested a sister group relationship with the small ternreks ( Microgale ; including the water ternreks ). Both authors also used skeletal anatomical features to check their DNA analyzes. However, RNA studies published three years earlier spoke out against being included in the travel treks; they placed the Erdtenrek at the base of the development of the Tenreks. Further DNA analyzes later confirmed that the Erdtenrek forms an independent subfamily, but compared these to the Reistenreks, which resulted in a total of three different phylogenetic positions. The differing genetic data suggested whether the genus Geogale is more diverse and possibly contains more than one species. The view is supported by a DNA study by a research team led by Kathryn M. Everson from 2016, in which all known representatives of the Tenreks were taken into account and several individuals of the Earth Tenrec were sequenced. In it, the authors conclude that the Erdtenrek actually includes several cryptic species , the separation of which already occurred in the Upper Miocene about 9 million years ago.

Subfossil finds

Some subfossil finds are documented from individual Malagasy caves . These include the Ankilitelo Cave , which is located on the southern edge of the Mikoboka Plateau in southwestern Madagascar. The cave has been scientifically researched since 1994 and contains a very rich fauna that includes half a dozen species of tenreks as well as various primates , bats and rodents . The age of the finds is around 510 to 630 years. Several partial skulls and jaw fragments were recovered from the Erdtenrek. Other remains of the Erdtenreks come from the Andrahomana Cave in the extreme southeast of Madagascar. A rich ensemble of finds is also documented from here, covering the period of around 8,700 years. The exploration of the Andrahomana Cave dates back to the end of the 19th century. As early as 1928, Guillaume Grandidier had referred a skull and lower jaw to the species Cryptogale australis , whereby the generic name refers to the cave site and the species name refers to the location of the cave very far in the south of Madagascar. Heim de Balsac then saw in 1972 Cryptogale australis as identical to the Erdtenrek due to the similar set of teeth and synonymized both forms. Both the Ankilitelo Cave and the Andrahomana Cave are in the distribution area of ​​the Erdtenreks, but the latter in the southeastern band area. The area around the Ankilitelo Cave is today characterized by a very dry climate, the Andrahomana Cave is located in the transition area of ​​the dry areas to the western one to the humid zones of eastern Madagascar.

Erdtenreks in Africa?

In 1957, Percy M. Butler and A. Tindell Hopwood presented fossilized tenre remains from East Africa . Among them were a part of the skull in the East African was Victoria nearby island of Rusinga . This was found in the Hiwegi Formation , which had formed in the Lower Miocene about 17.5 million years ago. A special feature of the skull was the set of teeth, which resembled those of the Erdtenreks, but was additionally reduced by the second premolar in the upper jaw. In its dimensions, the skull exceeded that of the Erdtenreks and reached about the size of the skull of the Cowan small tenreks ( Microgale cowani ). Butler and Hopwood referred the find to the species Geogale aletris . In the period that followed, another partial skull and a lower jaw fragment came to light from the Kapurtay agglomerates of Chamtwara in the Nyanza region of Kenya, which is about the same age, to the east of Lake Victoria . The inclusion of the fossil material in the current genus Geogale has been criticized several times, Butler himself created the generic name Parageogale in 1984 as an alternative . In doing so, he took into account some primeval features, mainly in the formation of the cusps on the molars, which connect the skull fragments with other fossil tenreks such as Protenrek , as well as the more reduced dentition compared to the earth tenrek . The latter rules out a direct precursor of parageogals to the Erdtenrek. The following year, Walter and Christl Poduschka suggested the alternative name Butleriella , which is now a synonym. Since there are no pre- Pleistocene tenreks fossils from Madagascar , the African finds are of great importance for the history of the origin and development of the family as well as for the settlement history of Madagascar and the later adaptive radiation there. Several phylogenetic studies confirm the closer relationships between geogals and parageogals .

Threat and protection

Due to the wide distribution and the assumed large population, the IUCN classifies the Erdtenrek as "not threatened" ( least concern ). The clearing of forests and conversion of the land into agricultural areas or the removal of trees for charcoal production pose greater threats to local populations . The Erdtenrek occurs in several protected areas, such as the Andohahela National Park , the Zombitse-Vohibasia National Park and the Tsimanampetsotsa National Park . It has also been found in some private protected areas such as the Kirindy forest area.

literature

  • JF Eisenberg and Edwin Gould: The Tenrecs: A Study in Mammalian Behavior and Evolution. Smithsonian Institution Press, 1970, pp. 1-138
  • 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
  • Nick Garbutt: Mammals of Madagascar. A complete guide. Yale University Press, 2007, pp. 1–304 (pp. 32–56)
  • Guillaume Grandidier and Gabriel Petit: Etude d'un mammifère insectivore malgache. Le Geogale aurita Alph. Milne-Edwards and Alfred Grandidier. In: A. Gruvel (Ed.): Faune des Colonies Françaises. Tome Quatrième. Paris, 1930, pp. 441–492 ( [9] )
  • 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 (pp. 165–166) ISBN 978-84-16728-08-4
  • Alphonse Milne-Edwards and Alfred Grandidier: Description d'un nouveau mammifère insectivore de Madagascar (Geogale aurita). Annales des Sciencas Naturelles, Zoologie 15, 1872, pp. 1–5 ( [10] )
  • Ronald M. Nowak: Walker's Mammals of the World. The Johns Hopkins University Press, Baltimore 1999, ISBN 0-8018-5789-9

Individual evidence

  1. a b c d Guillaume Grandidier and Gabriel Petit: Etude d'un mammifère insectivore malgache. Le Geogale aurita Alph. Milne-Edwards and Alfred Grandidier. In: A. Gruvel (Ed.): Faune des Colonies Françaises. Tome Quatrième. Paris, 1930, pp. 441-492
  2. ^ A b c Steven M. Goodman, Paulina D. Jenkins and Mark Pidgeon: Lipotyphla (Tenrecidae and Soricidae) of the Réserve Naturelle Intégrale d'Andohahela, Madagascar. Fieldiana Zoology 94, 1999, pp. 187-216
  3. a b c d e f g h i j Peter J. Stephenson: The Large-eared tenrec (Geogale aurita). Afrotherian Conservation 1, 2002, pp. 1-3
  4. a b c d e f Nick Garbutt: Mammals of Madagascar. A complete guide. Yale University Press, 2007, pp. 1–304 (pp. 32–56)
  5. ^ A b W. Poduschka and C. Poduschka: On the question of the generic name of “Geogale” aletris Butler and Hopwood, 1957 (Mammalia, Insectivora) from the Miocene of East Africa. Zeitschrift für Mammalskunde 50, 1985, pp. 129-140 ( [1] )
  6. a b 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 ( [2] )
  7. a b c d e f g h 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 (pp. 167–168) ISBN 978-84-16728-08-4
  8. ^ 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
  9. ^ 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
  10. ^ Justine A. Salton and Eric J. Sargis: Evolutionary Morphology of the Tenrecoidea (Mammalia) Hindlimb Skeleton. Journal of Morphology 270, 2009, pp. 367-387
  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. a b c d Voahangy Soarimalala: Les Afrosoricides de la forêt sèche malgache. Afrotherian Conservation 8, 2011, pp. 4-9
  13. ^ Daniel Rakotondravony, Steven M. Goodman, Jean-Marc Duplantier and Voahangy Soarimalala: Les petits mammifères. In: Joelisoa Ratsirarson and Steven M. Goodman (eds.): Inventaire biologique de la forêt littorale de Tampolo (Fenoarivo Atsinanana). Antananarivo, 1998, pp. 197-212
  14. ^ A b Steven M. Goodman, John C. Mittermeier, Juliot Ramamonjisoa and Lily-Arison Rene de Roland: The dietary habits of Barn Owls (Tyto alba) in the spiny bush of southwestern Madagascar. Malagasy Nature 8, 2014, pp. 67-72
  15. ^ 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
  16. ^ A b c Peter J. Stephenson: Reproductive biology of the Large-eared tenrec Geogale aurita (Insectivora: Tenrecidae). Mammalia 57 (4), 1993, pp. 553-563
  17. Steven M. Goodman and Voahangy Soarimalala: A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Forêt des Mikea of ​​southwestern Madagascar. Proceedings of the Biological Society of Washington 117 (3), 2004, pp. 251-265
  18. ^ A b P. J. Stephenson, Voahangy Soarimalala and Steven M. Goodman: Geogale aurita. The IUCN Red List of Threatened Species 2016. e.T9048A97188944 ( [3] ); last accessed on January 29, 2017
  19. ^ Edwin Gould and John F. Eisenberg: Notes on the biology of the Tenrecidae. Journal of Mammalogy 47 (4), 1966, pp. 660-686
  20. ^ A b c J. F. Eisenberg and Edwin Gould: The Tenrecs: A Study in Mammalian Behavior and Evolution. Smithsonian Institution Press, 1970, pp. 1-138
  21. ^ A b c Peter J. Stephenson, Paul A. Racey and Félix Rakotondraparany: Maintenance and reproduction of tenrecs (Tenrecidae) at Parc Tsimbazaza, Madagascar. International Zoo Yearbook 33, 1994, pp. 194-201
  22. Steven M. Goodman, Olivier Langrand and Christopher J. Raxworthy: Food habits of the Madagascar Long-eared owl Asio madagascariensis in two habitats in Southern Madagascar. Ostrich: Journal of African Ornithology 64 (2), 1993, pp. 79-85
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  26. a b c 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
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