Kleintenreks

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Kleintenreks
Systematics
Superordinate : Afrotheria
without rank: Afroinsectiphilia
Order : Tenrecus (Afrosoricida)
Family : Tenreks (Tenrecidae)
Subfamily : Travel treks (Oryzorictinae)
Genre : Kleintenreks
Scientific name
Microgale
Thomas , 1882

The Kleintenreks ( Microgale ), sometimes referred to as shrew or long-tailed treks, are a genus of mammals from the subfamily of Reistenreks (Oryzorictinae) within the family of Tenreks (Tenrecidae). They are small representatives of the Tenreks, which in their habitus resemble the shrewsresemble. Typical are the spindle-shaped body, the strong limbs and the long, narrow head. The fur has a soft texture, the tail varies noticeably in length depending on the species, with some it is only half as long as the rest of the body, with others it is twice as long.

The 21 known species of Kleintenreks are endemic to Madagascar , a larger part is adapted to the tropical rainforests in the east of the island state, a smaller part also inhabits the drier areas in the west. There is hardly any information about the way of life, the animals live hidden and are rarely seen. However, due to the varying appearance, caused by the different tail length and foot size, there are different types of adaptation, ranging from tree climbing, to living in the ground to digging underground. Only one form, the water tenrek , is adapted to life in water.

The genus was introduced in 1882, along with a long- and a short-tailed form. By the 1930s, almost two dozen species had been described, but their validity was discussed in detail. Overall, the information remained sparse and the number of known individuals was rather small. Only in the course of intensive research into the biological diversity of Madagascar, which began in the 1990s, did knowledge of the animal group increase significantly. Genetic examinations from 2016 took into account all known representatives for the first time and thus provided insight into the existing relationships.

features

Habitus

Kleintenreks are similar to the shrews - which do not exist in Madagascar - and apparently have occupied the same ecological niches as these. They are among the smaller representatives of tenrecs their head-body length ranges from 4.7 to 6.6cm in Zwergkleintenrek ( Microgale parvula ) and the Gnomkleintenrek ( Microgale pusilla ) to 12.2 to 17.0 cm for the web-footed tenrec ( Microgale mergulus ). The body weight therefore fluctuates from 2.1 to 4.2 g for the smallest and from 80 to 105 g for the largest forms. The tail is of different lengths in the individual species, it can only take up half the length of the rest of the body as in the short-tailed Kleintenrek or the Grandidier-Kleintenrek ( Microgale grandidieri ) or it can exceed the head-trunk length by more than twice , such as the small long-tailed small tenrek ( Microgale longicaudata ) and the large long-tailed small tenrek ( Microgale principula ). Expressed in absolute numbers, the tail length ranges from 3.3 to 17.1 cm. The body has a characteristic spindle-shaped structure, the limbs are short and strong. The muzzle is elongated, the ears protrude from the fur, and the long vibrissae are also striking . The fur has a soft texture, its color varies on the upper side from reddish brown through various brown and gray tones to almost black, the underside is usually lighter in color. The hands and feet each have five clawed rays, the three in the middle of which are mostly stronger. On the underside there are four skin pads, one each at the base of the first and fifth rays and one each at the base between the second and third and third and fourth rays. In addition, a thenar and a hypothenar are formed, but both have reduced sizes.

Skull and dentition features

The size of the skull varies greatly, the greatest length ranges from 15.5 to 32.0 mm in length, the greatest width on the cranium from 6.0 to 16.5 mm. Overall, it shows only a few changes within the genres. It is elongated and narrow, the rostrum tapers in most species. The extended posterior part of the skull appears conspicuous compared to the anterior part, the ratio of the Cowan-Kleintenrek ( Microgale cowani ) is 0.52: 0.48. In the case of the other tenreks, the relationship is reversed and is reinforced in the case of the hedgehog tenreks (Tenrecinae) by the extension of the rostrum. A characteristic feature is the non-closed zygomatic arch . In contrast to other tenreks, the pair of nasal bones is not fused, and there is no crest .

The Kleintenreks have only a slightly reduced number of teeth compared to the complete set of teeth of the higher mammals , only the foremost premolar has receded. The bit is thus composed of 40 teeth together and has the following dental formula : . The two front incisors in each case are always larger than the third, with the first incisor in the upper row of teeth and the second in the lower row often being the largest. The canine tooth is similar to the incisor teeth and is therefore not very specialized. The same can be said for the anterior premolars, which are mostly simply designed. The front teeth up to and including the front premolars often have additional small cusps in front of and behind the tooth crown, and depending on the type, they are separated from each other by diastemas of varying position and length. The posterior premolar resembles the subsequent molars , the latter showing only a few differences in the Kleintenreks. The molars are characterized by a zalambdodontic chewing surface pattern. It consists of three main tubercles (Para-, Proto- and Metaconus; related to the maxillary molars) in a triangular arrangement, whereby the Paraconus has a particularly large and pyramidal shape. The last upper molar is characteristically reduced in size, especially the posterior parts of the tooth.

Skeletal features

In the postcranial skeletal structure, the lower number of thoracic and lumbar vertebrae is striking, which together comprise 21 to 22 elements in the small tenreks compared to 22 to 23 in the closely related rice treks ( Oryzorictes ) or up to 24 in the more distantly related large tenreks ( Tenrec ). The number of tail vertebrae fluctuates greatly, depending on the length of the tail, it is composed of 24 vertebrae in the Cowan Kleintenrek, 32 in the Gnome Kleintenrek and 44 to 52 vertebrae in the Small Long-tailed Kleintenrek. In addition to the long-tailed pangolin ( Phataginus tetradactyla ), the latter have the highest number of individual vertebrae in the tail of today's higher mammals.

distribution

Kleintenreks are among the endemic inhabitants of Madagascar . They occur in a variety of different habitats , but usually rely on dense vegetation. A large part of the species is adapted to the tropical rainforests and sometimes also to the marshland of the eastern part of the island. With only a few exceptions, these show an extensive distribution area that, often oriented north-south, extends in a more or less wide strip over almost the entire length of the island of 1300 km in length. What is noticeable about these species is the wide range of elevations, ranging from the lower and middle to the higher mountain levels around 2500 m, such as the Drouhard-Kleintenrek ( Microgale drouhardi ), which can occur at heights of 360 to 2350 m, or the Dwarf lesser rek ( Microgale parvula ), which occurs between 450 and 2050 m altitude. A smaller part of the species has a limited range both geographically and in terms of altitude, here the mountain small tenrek ( Microgale monticola ) belongs to it. Less than a quarter of the representatives populate the drier landscapes in the west, the individual occurrences are spatially limited with the exception of the short-tailed small ternrek ( Microgale brevicaudata ). It is particularly noticeable that in the mountainous landscapes of eastern Madagascar numerous species of Kleintenreks appear more or less sympatric . A total of a dozen species have been recorded on the Tsaratanana massif in the province of Mahajanga , in the forest area of ​​Tsinjoarivo in the central Malagasy province of Antananarivo there are just as many, while in several other regions up to ten different species can be found. In principle, the different niche uses of the Kleintenreks are largely unexplored. In individual regions, deviations in the height distribution of individual species were found, for example in the small ( Microgale longicaudata ) and the large long-tailed Kleintenrek ( Microgale principula ) on the Anjanaharibe massif in the Antsiranana province , but this could not be confirmed in other areas examined.

Way of life

General

The way of life of the Kleintenreks has only been little studied, they tend to live hidden away and are rarely seen in the wild. The few better studied species include the water tenrek ( Microgale mergulus ) and the Cowan small tenrek ( Microgale cowani ). The small tenreks are land-dwelling with the exception of the water tenreks, which are the only representatives of the tenreks that are adapted to a semi-aquatic way of life. In general, the assessments of the behavior of the other species are based on the external characteristics, less often on actual observations. Thus, for the small ( Microgale longicaudata ) and the large long-tailed small tenrek ( Microgale principula ) or the major long-tailed small tenrek ( Microgale majori ), a partially tree-climbing locomotion is assumed, to which they are adapted with a long, flexible tail and elongated feet . The long tail then functions as a grasping organ , which has been proven at least in the small long-tailed small tenrek. The rather short-tailed species such as the short-tailed Kleintenrek ( Microgale brevicaudata ), the bare-nosed Kleintenrek ( Microgale gymnorhyncha ) or the Delicate Kleintenrek ( Microgale gracilis ), on the other hand, are probably partly underground burrowing forms. This is also supported by the rather broad feet with long claws in these species, as well as the small eyes and also small ears that are almost hidden under the fur. Animals with intermediate tails that reach about the length of the rest of the body, on the other hand, are considered to be ground dwellers, such as the Cowan-Kleintenrek ( Microgale cowani ), the Thomas-Kleintenrek ( Microgale thomasi ) or the Taiva-Kleintenrek ( Microgale taiva ). Hardly any information is available about the animals' activity times, and some species such as the small long-tailed small tenrek are nocturnal. Sometimes animals also create nests as protection and shelter.

Diet and energy balance

Information on the diet of the small tenreks is only sparingly available; in general, as with most of the other relatives, a carnivorous to insectivorous diet (faunivor) can be assumed. According to examinations of excrement residues from the Ranomafana forest area, the water tenrek mainly preyed on insects , occasionally also crabs or young frogs . Something similar is known of the Cowan-Kleintenrek and the small long-tailed Kleintenrek. Isotope studies on a dozen different species of small tenreks from the Tsinjoarivo forest area confirm the largely faunivorous way of life. But they also reveal individual differences between the species. The gnome lesser rek ( Microgale pusilla ) with its high nitrogen content differs significantly from the long-tailed lesser rek with significantly lower values. For the former, a preference for predatory prey is to be assumed, for the latter it is more likely for herbivorous animals. The strongly varying carbon values ​​in the various representatives of the long-tailed small tenreks also suggest that food is sought in different levels of the forests. In contrast, burrowing species show a very narrow and higher range of nitrogen values ​​and thus stand out from their tree relatives. The investigations thus give a first insight into the trophic differentiation of the small tenreks in a narrowly limited and largely shared habitat .

The average body temperature is 32 to 34 ° C. Unlike other Tenreks, it is largely stable, but the information is based on only three investigated species. The resting metabolic rate varies between 76 and 138% of the expected value for mammals of the same size. Among other things, it increases in the reproductive phase. The values ​​for both body temperature and metabolism are on average higher than those of the close relatives of the genus Nesogale .

Reproduction

The reproduction is hardly documented, one litter has so far only been observed in the Cowan-Kleintenrek. This comprised three young animals, each weighing around 2.5 g. As is customary with Tenreks, the newborns were born as nestlings with bare skin and closed ears and eyes. They only survived a few days, so nothing is known about their further development. In the wild, pregnant or milk-producing females were mostly observed in the period from October to December, as well as males with enlarged testicles and animals with pronounced milk teeth . No information is available on life expectancy in the wild, most animals in captivity have not survived longer than two months, and one individual of the Cowan Lesser Tenderen spent more than a year in human care.

Systematics

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 Kleintenreks are a genus within the Tenreks family (Tenrecidae). Together with the rice burrowers ( Oryzorictes ) and the members of the genus Nesogale, they form the Reistenreks (Oryzorictinae), one of the three subfamilies of the Tenreks. The other two are represented by the spiky hedgehog tenreks (Tenrecinae) and the monotypical earth tenreks (Geogalinae). With more than 20 species, the Kleintenreks form the most varied member of the family and make up around two thirds of today's representatives of the Tenreks. Due to some morphological features, they are considered to be more originally within the family, including the barely reduced set of teeth with the poorly differentiated canine, the small-scale specialized skull structure compared to the long-snouted hedgehog tenreks, the rather low number of thoracic and lumbar vertebrae in comparison the higher number in some other tenreks and the formation of a soft, unspecialized fur in contrast to the spines of the hedgehog tenreks. According to molecular genetic analyzes, the genus originated in the Lower Miocene around 16.8 million years ago and diversified very strongly in the period that followed. The speciation rate was significantly higher in the moist tropical forests than in the drier parts of the island. The separation from the common ancestor with the genus Nesogale as the closest relative of the Kleintenreks began 19.4 million years ago.

There are currently 21 different types:

Internal systematics of the Kleintenreks according to Everson et al. 2016
 Microgale  



 Microgale pusilla


   

 Microgale majori


   

 Microgale principula


   

 Microgale jenkinsae


   

 Microgale longicaudata






   

 Microgale mergulus


   

 Microgale parvula




   



 Microgale brevicaudata


   

 Microgale grandidieri



   

 Microgale drouhardi


   

 Microgale monticola


   

 Microgale taiva





   



 Microgale gracilis


   

 Microgale thomasi


   

 Microgale cowani


   

 Microgale jobihely





   

 Microgale dryas


   

 Microgale gymnorhyncha




   

 Microgale soricoides


   

 Microgale fotsifotsy


   

 Microgale nasoloi







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Originally, the Dobsonian Kleintenrek ( Nesogale dobsoni ) and the Talazac-Kleintenrek ( Nesogale talazaci ) were also members of the genus Microgale , but today they are assigned to Nesogale . The water tenrek, on the other hand, was part of its own genus Limnogale for a long time . The partially independent form Microgale prolixacaudata , currently a synonym of the small long-tailed small tiger ( Microgale longicaudata ), has not yet been genetically investigated in more detail. On the basis of haplotypes , however, a narrowly outlined clade , restricted to northern Madagascar, can be separated from the small long-tailed small tenrek, which possibly corresponds to this shape.

In addition to the species that exist today, there is also an extinct representative, which has been documented using subfossil material:

Other subfossil forms were introduced with Microgale breviceps and Microgale decaryi . The former is now considered to be identical to the short-tailed small tenrek, the latter is a synonym for the large long-tailed small tenrek.

Research history

On the genus history of Microgale

Oldfield Thomas

The Kleintenreks as a generic form were established by Oldfield Thomas in 1882 . At that time Thomas examined various individuals from the forest area of ​​Ankafina around 10 km south of Ambohimahasia in central-eastern Madagascar, which had already been picked up by W. Deans Cowan two years earlier. In his first description , Thomas made a distinction between the small long-tailed Kleintenrek and the Cowan-Kleintenrek, two types of which the former represents the type form. He saw the genus Microgale at this time as a link between the tenreks and the otter shrews , with the tenreks they shared the formation of a collarbone and a teardrop canal (Canalis lacrimale), with the otter shrews the long tail, the soft fur and the tibia and fibula, which are fused together in the lower half . The name Microgale is derived from the Greek words μικρός ( mikrós ) for "small" and γαλἑη ( gale ) for "weasel"; the ending gale is often found in connection with tenre kites or other insectivorous animals.

In 1918, Oldfield Thomas caused the relatively high number of around a dozen species names assigned to the Kleintenreks to reorganize the group at genus level. He split up Microgale and united the Dobsonian Kleintenrek and the Talazac Kleintenrek in the genus Nesogale . Both species have an average larger body shape than most other small tenreks (the water tenrek is an exception), plus the very robust skull with prominent muscle marks. Terence Morrison-Scott confirmed the independence of Nesogale in 1948 , but subsequent authors did not recognize it, such as Henri Heim de Balsac in 1972, who formally abolished the genre and reunited Nesogale with Microgale . Ross DE MacPhee carried out a comprehensive revision of the Kleintenreks in 1987 and underpinned it with morphological and morphometric investigations. He largely followed Heim de Balsac's view, especially since he could hardly work out any deviating characteristics. The status quo remained then obtained by this revision for the next nearly three decades. Molecular genetic analyzes, which were published in 2016, revealed a very early separation of the Dobsonian and Talazac lesser tenreks from the other lesser tenreks, which prompted the authors to reassign the genus status to Nesogale and to incorporate the two species there.

Charles Immanuel Forsyth Major described the water tenrek in 1896 and classified it in its own genus Limnogale , which he defined via the trained webbed feet, the powerful tail and individual features of the skull and teeth. The genus lasted for more than a hundred years and was hardly doubted; some researchers pointed out the existing similarities to the Kleintenreks. The exact phylogenetic position was the subject of discussion and varied between the sister form of the otter shrew and that of the tenreks. In the first more comprehensive genetic studies published in 2003, Limnogale was found to be deeply embedded in the genus Microgale , making the Kleintenreks a paraphyletic group. Since the result was confirmed in later analyzes, the water tendril was transferred to Microgale in the course of the latest genetic investigations from 2016 and the genus Limnogale was dissolved.

Together with Nesogale , Thomas had also introduced Leptogale as a further genus and assigned it to the graceful Kleintenrek, whose skull is characteristically slim and has an elongated rostrum . Thirteen years later established Guillaume Grandidier and Gabriel Petit with Paramicrogale another genre whose hallmark was a robust skull. The authors referred to the genus, among other things, Paramicrogale occidentalis , a species that is not valid today, which is identical to the short-tailed Kleintenrek. In the period that followed, however, both genres were only rarely recognized or, in some cases, only classified at sub-genre level. In his revision, MacPhee rated both as synonymous with Microgale .

Attempts at structuring the Kleintenreks

Already in the first description of the genus Microgale , Thomas pointed out the striking differences in tail length and the shape of the foot in the two species he presented. By the mid-1930s, more than 20 species had already been introduced, which were quite varied in terms of their external characteristics. At this time, Guillaume Grandidier used the tail length for the first time for a more precise structure of the small tenreks. He distinguished three groups of forms:

  • long-tailed animals (with a tail twice as long as the rest of the body): M. longicaudata , M. majori , M. principula (including M. sorella )
  • short-tailed animals (with a tail that is shorter than the rest of the body): M. brevicaudata , M. cowani (including M. crassipes , M. longirostris , M. nigrescens ), M. thomasi
  • intermediate forms (with a tail about as long as the rest of the body): M. drouhardi , M. parvula , M. pusilla , M. taiva

Terence Morrison-Scott took up Grandidier's attempt at structuring later, but reduced the number of species through individual synonymizations.

In the 1970s, John F. Eisenberg and Edwin Gould refined Grandidier's approaches and began to classify the large number of species according to functional morphological aspects. The ratio of the length of the tail or the size of the foot to the length of the head and trunk was taken as the basis. A short tail is associated with a fossorial (digging), a very long one with an arboreal (tree-climbing) or scansorial (sometimes tree-climbing) way of life. The size of the foot, on the other hand, provides information as to whether an animal moves on two-legged hopping ( saltatory ; with large feet) or four-footed (with smaller feet). As a result, Eisenberg and Gould differentiated a total of four eco-morphotypes in 1970:

  • Forms that sometimes live underground and have little ability to jump (ratio of tail to body length 0.5: 1, ratio of hind foot to body length less than 0.2: 1): M. brevicaudata
  • Forms that search for food on the surface of the earth and may have a certain ability to climb or jump (ratio of tail to body length 0.6–1: 1, ratio of hind foot to body length 0.21–0.28: 1): M. cowani (including M. longirostris ), M. gracilis , M. occidentalis (= M. brevicaudata ), M. thomasi , Nesogale dobsoni
  • Forms that search for food and can climb on the surface of the earth (ratio of tail to body length 1–1.5: 1, ratio of hind foot to body length 0.2–0.27: 1): M. drouhardi (including M. melanorrhachis ), M. parvula , M. pusilla , Nesogale talazaci
  • Forms that can climb and possibly live in the branches (ratio of tail to body length 1.5–2.6: 1, ratio of hind foot to body length greater than 0.26: 1): M. longicaudata (including M. prolixacaudata ), M. majori , M. principula (including M. sorella )

In the above-mentioned revision of the Kleintenreks by Ross DE MacPhee from 1987, he included a large part of the specimens of the genus kept in museums and thus known at that time. The number studied comprised about 120 individuals. Most of the studies on the Kleintenreks were hardly based on statistical analyzes until then, rather they were based on more or less empirical data. MacPhee thus created a morphological and morphometric database for the first time. This led to the fact that he considered some of the forms described as synonyms and thus reduced the number of species from over twenty to ten. In a further step, he combined these into groups, which he called clusters . The individual clusters represented less of an actual family group, rather they expressed similarities in the phenotype :

  • cowani clusters: M. cowani , M. thomasi , M. parvula
  • gracilis cluster: M. gracilis
  • longicaudata cluster: M. longicaudata , M. principula
  • pusilla cluster: M. pusilla
  • brevicaudata cluster: M. brevicaudata
  • dobsoni clusters: Nesogale dobsoni , Nesogale talazaci

The division into different clusters was later taken up and expanded by some authors. So led Paulina D. Jenkins in 1996 the soricoides clusters based on the Shrew Kleintenrek and expanded a variety of other clusters to new or re-recognized species.

MacPhee's detailed work turned out to be problematic in some cases, as some of the species described up to 1987 were only known through the respective type specimens. In these cases, individual synonymizations were based on too little data, so that no sufficient differentiation of characteristics could be made. In the 1990s, an intensive phase of research into the biological diversity of Madagascar began, during which numerous natural areas on the island were inventoried and cataloged. For some species, including synonymized forms with mostly singular evidence, this led to the discovery and provision of new comparative material and thus to the confirmation or recognition of the independent species status, for example in the major long-tailed small tenrek, the Taiva small tenrek or the Drouhard Kleintenrek. In addition, numerous new species could be discovered and described, altogether this affects almost half of the small tenre maps known today. As a result of this research activity, the number of recognized representatives of the Kleintenreks increased again to a total of 21 by 2009 (the year of the last new species description).

The emergence of molecular genetic investigation procedures in the transition from the 20th to the 21st century made it possible not only to determine the phylogenetic position of the Tenreks, and thus also of the Kleintenreks, more precisely, but also to shed light on the internal relationship of the individual groups. This resulted very early on in the fact that the water tenrek, which was not taken into account in the previous morphological and morphometric investigations, was recognized as a clear representative of the small tenreks. The analyzes from 2016 are of importance here, which for the first time included all species of Tenreks known at that time and grouped the individual representatives conclusively. The rather unexpected results showed on the one hand the sister group ratio of the smallest (dwarf lesser trek) and the largest (water tenrek) representative of the lesser tenreks, and on the other hand that the development of the four species adapted to dry climates (short-tailed, nasolo and Jenkins lesser treks as well as grandidians) -Kleintenrek) can be traced back to a total of three independent events. However, further studies of the biodiversity of Madagascar are necessary to clarify such processes more precisely.

Tribal history

Fossil finds of Kleintenreks from past geological periods are not available, but there are some subfossil remains from the Holocene . They come mainly from two important cave sites. One of them is the Andrahomana Cave southwest of Tolagnaro in the extreme southeast of Madagascar, whose rich bone and tooth material covers the period of the last 8700 years. Among the numerous mammals there are also the remains of tenre karts. These include four species of Kleintenreks still alive today: the small and large long-tailed Kleintenrek, the Gnome Kleintenrek and the Nasolo Kleintenrek. In addition, the extinct form Microgale macpheei occurs, the remains of which date back to between 1600 and 2700 years. The second important site is in the Ankilitelo Cave on the southern edge of the Mikoboka Plateau in southwestern Madagascar. Here, too, there is an extensive fauna with at least three species of small tenrek, in addition to the nasolo small tenrek, the short-tailed small tenrek and the major long-tailed small tenrek have been detected. The finds are between 510 and 630 years old. Both regions are characterized by dry climatic conditions and are outside the range of the recent representatives. The disappearance of the various species may be related to the increasingly dry climate in western Madagascar, but at that time humans were already populating the island. There are also a few other caves or sites with subfossil remains, for example near Mahajanga in northwestern Mozambique, or in the area of ​​the Tsimanampetsotsa National Park in the southwestern part of the island.

Threat and protection

Due to the loss of habitat that lists IUCN six species as "critically endangered" ( endangered ) or "at risk" ( vulnerable ). In the former category belong the Jenkins-Kleintenrek ( Microgale jenkinsae ) and the Dunkle Kleintenrek ( Microgale jobihely ), in the latter the Dryad-Kleintenrek ( Microgale dryas ), the Water-Ternrek ( Microgale mergulus ), the Mountain-Kleintenrek ( Microgale monticola ) and the Nasolo -Kleintenrek ( Microgale nasoloi ). The species mentioned have only a limited range, all other small tenreks occur more or less over large parts of Madagascar. The main threats to the existence of individual populations are the destruction of forests through logging, conversion into agricultural land and fire; locally, mining also has a certain influence. With a few exceptions, all species are present in nature reserves; field research into their biology and ecology is necessary to further protect the small tenreks.

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
  • 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
  • RDE MacPhee: The Shrew Tenrecs of Madagascar: Systematic Revision and Holocene Distribution of Microgale (Tenrecidae, Insectivora). American Museum Novitates 2889, 1987, pp. 1-45
  • 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 c 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
  2. ↑ a b c 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
  3. ↑ a b c Link E. Olson, Z. Rakotomalala, KBP Hildebrandt, HC Lanier, Christopher J. Raxworthy and Steven M. Goodman: Phylogeography of Microgale brevicaudata (Tenrecidae) and description of a new species from Western Madagascar. Journal of Mammalogy 90 (5), 2009, pp. 1095-1110
  4. ↑ a b c d e Steven M. Goodman and Paulina D. Jenkins: The Insectivores of the Réserve Spéciale d'Anjanaharibe-Sud, Madagascar. Fieldiana Zoology 90, 1998, pp. 139-161
  5. ^ 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
  6. ↑ a b c d Steven M. Goodman and Paulina D. Jenkins: Tenrecs (Lipotyphla; Tenrecidae) of the Parc National de Marojejy, Madagascar. Fieldiana Zoology 97, 2000, pp. 201-229
  7. ↑ a b c d e f g R. DE MacPhee: The Shrew Tenrecs of Madagascar: Systematic Revision and Holocene Distribution of Microgale (Tenrecidae, Insectivora). American Museum Novitates 2889, 1987, pp. 1-45
  8. ^ A b Steven M. Goodman, Christopher J. Raxworthy, CP Maminirina and Link E. Olson: A new species of shrew tenrec (Microgale jobihely) from northern Madagascar. Journal of Zoology 270, 2006, pp. 384-398
  9. ↑ Voahangy Soarimalala, Martin Raheriarisena and Steven M. Goodman: New distributional records from central-eastern Madagascar and patterns of morphological variation in the endangered shrew tenrec Microgale jobihely (Afrosoricida: Tenrecidae). Mammalia 74, 2010, pp. 187-198
  10. ↑ a b c d 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] )
  11. ^ A b c William King Gregory: The orders of mammals. Bulletin of the American Museum of Natural History 27, 1910, pp. 1–524 (pp. 236–240)
  12. ↑ a b c Oldfield Thomas: Description of a new genus and two new species of Insectivora from Madagascar. Journal of the Linnean Society of London 16, 1882, pp. 319–322 ( [2] )
  13. ↑ a b c Link E. Olson: Tenrecs. Current Biology 23 (1), 2013, pp. R5-R8
  14. ↑ Paulina D. Jenkins Christopher J. Raxworthy and Ronald A. Nussbaum: A new species of Microgale (Insectivora, Tenrecidae), with comments on the status of four other taxa of shrew tenrecs. Bulletin of the Natural History Museum of London (Zoology) 63 (1), 1997, pp. 1-12
  15. ↑ 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
  16. ↑ Steven M. Goodman, Daniel Rakotondravony, Marie Jeanne Raherilalao, Domoina Rakotomalala, Achille P. Raselimanana, Voahangy Soarimalala, Jean-Marc Duplantier, Jean-Bernard Duchemin and J. Rafanomezantsoa: Inventaire biologique de la Foret de Tsinjoarivo, Ambatolampy. Akon'ny Ala 27, 2000, pp. 18-27
  17. ^ A b Paulina D. Jenkins, Steven M. Goodman and Christopher J. Raxworthy: The Shrew Tenrecs (Microgale) (Insectivora: Tenrecidae) of the Réserve Naturelle Intégrale d'Andringitra, Madagascar. Fieldiana Zoology 85, 1996, pp. 191-217
  18. ↑ Voahangy Soarimalala and Martin Raheriarisena: The non-volant and non-primate mammals of the Ambatovy Analamay forest. In: Steven. M. Goodman and V. Mass (Eds.): Biodiversity, exploration, and conservation of the natural habitats associated with the Ambatovy project. Malagasy Nature 3, 2010, pp. 153-177
  19. ↑ a b c d e f J. F. Eisenberg and Edwin Gould: The Tenrecs: A Study in Mammalian Behavior and Evolution. Smithsonian Institution Press, 1970, pp. 1-138
  20. ↑ 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
  21. ↑ a b Roland D. Hilgartner: Some ecological and behavioral notes on the shrew tenrec Microgale cf. longicaudatain the dry deciduous forest of western Madagascar. Afrotherian Conservation 3, 2005, pp. 3-5
  22. ^ 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
  23. ^ Paul A. Racey and Peter J. Stephenson: Reproductive and energetic differention of the Tenrecidae of Madagascar. In: WR Lourenço (ed.): Biogéographie de Madagascar. Paris, 1996, pp. 307-319
  24. ↑ Peter J. Stephenson: Resting metabolic rate and body temperature in the aquatic tenrec Limnogale mergulus (Insectivora: Tenrecidae). Acta Theriologica 39 (1), 1994, pp. 89-92
  25. ^ A b 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
  26. ↑ a b c d e f g h 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
  27. ↑ 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
  28. ↑ a b Link E. Olson, Steven M. Goodman and Anne D. Yoder: Illumination of cryptic species boundaries in long-tailed shrew tenrecs (Mammalia: Tenrecidae; Microgale), with new insights into geographic variation and distributional constraints. Biological Journal of the Linnean Society 83, 2004, pp. 1-22
  29. ↑ a b Martin Nicoll and Nanie Ratsifandrihamanana: The growth of Madagascar's protected areas system and its implications for tenrecs (Afrosoricida, Tenrecidae). Afrotherian Conservation 10, 2014, pp. 4-8
  30. ↑ a b IUCN: The IUCN Red List of Threatened Species. Version 2016.2. ( [3] ); last accessed on November 4, 2016
  31. ^ A b c Steven M. Goodman, Natalie Vasey and David A. Burney: Description of a new species of subfossil shrew tenrec (Afrosoricida: Tenrecidae: Microgale) from cave deposits in southeastern Madagascar. Proceedings of the Biological Society of Washington 120 (4), 2007, pp. 367-376
  32. ^ A b Walter Kaudern: Quaternary fossils from Madagascar. Zoological Yearbooks 41, 1918, pp. 521–534 ( [4] )
  33. ↑ Guillaume Grandidier: Description de deux nouveaux mammifères insectivores de Madagascar. Bulletin du Muséum national d'histoire naturelle 34, 1928, pp. 63–70 ( [5] )
  34. ↑ Martin Pickford: Late Eocene Potamogalidae and Tenrecidae (Mammalia) from the Sperrgebiet, Namibia. Coomunications of the Geological Survey of Namibia 16, 2015, pp. 114–152
  35. ↑ a b Oldfield Thomas: On the arrangement of the small Tenrecidae hitherto referred to Oryzorictes and Microgale. Annals and magazine of natural history 14, 1918, pp. 302–307 ( [6] )
  36. ^ A b T. CS Morrison-Scott: The Insectivorous Genera Microgale and Nesogale (Madagascar). Proceedings of the Zoological Society of London 118, 1948, pp. 817-822
  37. ^ A b c Henri Heim de Balsac: Insectivores. In: R. Battistini and G. Richard-Vindard (eds.): Biogeography and ecology in Madagascar. The Hague, 1972, pp. 629-660
  38. ^ CI Forsyth Major: Diagnoses of new mammals from Madagascar. Journal of Natural History 18 (106), 1896, pp. 318–321 ( [7] )
  39. ↑ Christian Guth, Henry Heim de Balsac and M. Lamotte: Recherches sur la morphologie de Micropotamogale lamottei et l'évolution des Potamogalinae: écologie, denture, anatomie crânienne. Mammalia 23, 1959, pp. 423-447
  40. ^ Robert J. Asher: A morphological basis for assessing the phylogeny of the "Tenrecoidea" (Mammalia, Lipotyphla). Cladistics 15, 1999, pp. 231-252
  41. ↑ a b c Link E. Olson and Steven M. Goodman: Phylogeny and biogeography of tenrecs. In: Steven M. Goodman and Jonathan P. Benstead (Eds.): The natural history of Madagascar. University of Chicago Press, 2003, pp. 1235-1242
  42. ^ A b Matjaž Kuntner, Laura J. May-Collado and Ingi Agnarsson: Phylogeny and conservation priorities of afrotherian mammals (Afrotheria, Mammalia). Zoologica Scripta 40 (1), 2011, pp. 1-15
  43. ↑ Guillaume Grandidier and G. Petit: Un type nouveaux de centetide malgache, Paramicrogale occidentalis. Bulletin de la Société zoologique de France 56, 1931, pp. 126-139 ( [8] )
  44. ↑ Guillaume Grandidier: Deux nouveaux mammifère insectivores de Madagascar Microgale drouhardi et M. parvula. Bulletin du Muséum national d'histoire naturelle 2, 1934, pp. 474–476 ( [9] )
  45. ^ Paulina D. Jenkins and Steven M. Goodman: A new species of Microgale (Lipotyphla: Tenrecidae) from isolated forest in southwestern Madagascar. Bulletin of the Natural History Museum of London (Zoology) 65 (2), 1999, pp. 155-164
  46. ^ DA Burney, N. Vasey, LR Godfrey, Ramilisonina, WL Jungers, M. Ramarolahy and L. Raharivony: New Findings at Andrahomana Cave, Southeastern Madagascar. Journal of Cave and Karst Studies 70 (1), 2008, pp. 13-24
  47. ↑ Kathleen M. Muldoon, Donald D. de Blieux, Elwyn L. Simons and Prithijit S. Chatrath: The Subfossil Occurrence and Paleoecological Significance of Small Mammals at Ankilitelo Cave, Southwestern Madagascar. Journal of Mammalogy 90 (5), 2009, pp. 1111-1131
  48. ^ A b Steven M. Goodman and William L. Jungers: Extinct Madagascar. Picturing the island's past. University of Chicago Press, 2014, pp. 1–206 (pp. 65–73 and 94–101)

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